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VOL 96, PART 1 29 FEBRUARY, 1972 


TRANSACTIONS OF THE 


ROYAL SOCIETY 
OF SOUTH AUSTRALIA 


INCORPORATED 


CS eT ee 


CONTENTS 


Kott, Patricia’ The Ascidians of South Australia I. ge Gulf, St. Vincent 
Gulf and Encounter Bay - - 1 


Harris, Wayne K. New form species of pollen from southern Australian early 
Tertiary sediments - - - - - - - - 53 


Sa a a I ST SRE EIS RS 


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. 
VOLUME 96, 1972 
PART 1, 29 FEBRUARY 


Kott, Patricia The Ascidians of South Australia I. BESESETES Gulf, St. Vincent 
Gulf and Encounter Bay - . x 4 2 a 


Harris, Wayne K. New form species of pollen from southern Australian early 
Tertiary sediments - - - : - = = s 
PART 2, 31 MAY 


Preiss, W. VY. The systematics of South Australian Precambrian and Cambrian 
Stromatolites. Part 1 - - - - = 2 f 4 


Mawson, Patricia M. The nematode genus Maxvachonia (Oxyurata: Cosmo- 
cercidae) in Australian reptiles and frogs - - - - 


Mawson, Patricia M. Three new species of the genus Cloacina Linstow (Nema- 
toda: Strongylata) from macropod marsupials - - - 


Schodde, R., Mason, I., & Wolfe, T. O. Further records of the Pitted-shelled 
Turtle (Carettochelys insculpta) from Australia - - - 


Wollaston, Elise M. The morphology and relationships of Muellerena wattsti 
(Harvey) Schmitz (Ceramiaceae: Rhodophyta) - - - 
PART 3, 31 AUGUST 


Smith, Meredith J. Small fossil vertebrates from Victoria Cave, Naracoorte, 
South Australia. Il. Peramelidae, Thylacinidae and Dasyuridae 
(Marsupialia) - - - - - - - - - 


Mawson, Patricia M. The genus Acuwaria Bremser (Nematoda: Spirurida) in 
Australia - - - - - = = = = > 


Milnes, A. R., & Bourman, R. P. A Late Palaeozoic glaciated granite surface at 
Port Elliot, South Australia - a - . 3 : * 


Tyler, M. J., & Parker F. Additions to the hylid frog fauna of New Guinea, with 
description of a new species, Litoria timida - - - - 
PART 4, 30 NOVEMBER 


Kott, Patricia The Ascidians of South Australia II. Eastern Sector of The Great 
Australian Bight and Investigator Strait - - & + x 


Dubois, Georges, & Angel, L. Madeline Strigeata (Trematoda) of Australian Birds 
and Mammals from the Helminthological Collection of the Uni- 
versity of Adelaide - - +! - - - - - 


OBITUARY: KEITH RODNEY MILES - = B 2 ¢ 2 2 
Annual Report of Council, 1971-72 - = = 2 4 s - 3 
Award of the Sir Joseph Verco Medal - - = 7 > 4 a = 
Balance Sheet - - - - - 2 - : : = 2 5 


List of Fellows - : = 2 “ 2 3 = x m 4 ~/ 


53 


67 


101 


100 


115 


119 


125 


139 


149 


157 


165 


197 
216 
220 
221 
222 
223 


THE ASCIDIANS OF SOUTH AUSTRALIA I. SPENCER GULF, ST. 
VINCENT GULF AND ENCOUNTER BAY 


BY PATRICIA KOTT 


Summary 


A large and representative collection of Ascidiacea from St. Vincent Gulf and adjacent locations is 
discussed. Fifty-nine species are represented, of which Pyura scoresbiensis and Ctenicella antipoda 
are new to science. Ascidia aclara Kott, previously known from other Australian locations, and 
Aplitiium colelloides Herdman, previously known only from South Africa, are recorded from the 
area for the first time. 

The fauna of St. Vincent Gulf is typically of the Flindersian marine biogeographic region, but 
includes several endemic species. Morphological characteristics accounting for the success of 
certain species and groups of species sharing a habitat are indicated. 


THE ASCIDIANS OF SOUTH AUSTRALIA IL 
SPENCER GULF, ST. VINCENT GULF AND ENCOUNTER BAY 


by Patricia Kott* 


Sutmnmary 


A large and representative collection of Ascidiacea from St. Vincent Gulf and adjacent locitions 
is discussed. Fifty-nine species are represented, of which Pyyra scoreshiensix and Crenicella antipoda 


are new to science. 


Ascidia aclara Kott, previously known from other Australian locations, and 


Aplidium colelloides Herdman, previously known only from South Africa, are recorded from the are 


for the first time, 


The fauna of St. Vincent Gulf is typically of the Flindersian marine biogeographic region, but 


includes several endemic species, 


Morphological characteristics accounting for the success of certain 


species and groups of specics sharing a habitat are indicated. 


Jntroduction 


This large collection of ascidians, mainly 
from St. Vincent Gulf, South Australia, was 
made by Mr. 8S. A. Shepherd of the Depart- 
ment of Fisheries and Fauna Conservation, 
South Australia. It is a valuable and represen- 
tative collection and demonstrates the value of 
SCUBA collections of this benthic group from 
otherwise inaccessible localities. Colour notes 
made by the collector provide most useful data 
for comparison with the preserved specimens 
in which colours are generally lost or change 
completely. The large number of individuals 
of most species that are available in the col- 
lection has demonstrated a wide variability in 
certain characters and some synonymy has 
been established. 

Information on the environmental conditions 
operating in various locations, also supplied by 
the collector, has been related to the morpho- 
logy of the species prescnt to contribute to an 
assessment of selective mechanisms affecting 
the ascidians. Full station lists of species are 
also given to facilitate consideration of the 
faunal associations and their ecological rela- 
tionships. 

The specimens are deposited in the South 
Australian Muscum. 

The following species have previously been 
recorded from South Australia (Kott 1952, 
1957a, 1962, 1963) but were not in the present 
collection. 


Polyclinum nepiunium 

Polyclinum marsupiale 

A plidium flavolineatum 

A plidium australiensis 

Lissoclinym ostrearium 

Dideninum turritum 

Didemnurm angusti 

Didemnum pseudediplosama 

Trididenmum nataletise 

Trididemnum cerebrijorme 

Leptaclinides imperfectus 

Syniplewma viride 

Styela lohata 

Asterocarpa cerea 

Pyura stolonifera 

Zoogeography 

The fauna is typically that of the Flindersian 
marine region, together with Distaplia viridis 
which is also recorded from Part Phillip Bay, 
Ascidia aclara which has been taken from simi- 
lar sheltered locations on the Victorian, New 
South Wales, and Queensland coasts, and 
Aplidium colelloides, previously recorded from 
South Africa. The new species, Pyura scores- 
hiensis and Ctenicella antipoda, may be 
endemic. 

The records of A. colelloides from off South 
Africa and South Australia suggest a circum- 
polar distribution, as demonstrated for many 
ascidian species (Kott 1971a). A wide dis- 
persal of larvae, however, does not provide a 
satisfactory explanation for this pattern of dis- 


* Zoology Department, University of Queensland, Qld. Australia 4067. 


Trans. R. Soc. S. Aust. 96, Part 1, 29 February 1972, 


4 


tribution since, for successful sexual reproduc- 
tian, minimal population densities of adults are 
required. The existence of so many circum- 
polar species in the extant fauna may be the 
result of a slow rate of evolution and the per- 
sistence of relict forms in certain areas. 


Habit of the Ascidian Fauna 

In the present collection, ascidians have been 
taken frum a wide variety of locations, espe- 
cially in St, Vincent Gulf. The terminology 
qualifying the conditions encountered is partly 
that described by Shepherd & Womersley (1970) 
and Womersley & Edmonds (1958), as fol- 
lows: 

(1) “Reugh Coast Subformation” (R.C.S.) 

refers to coasts exposed to the southern 

ocean swell (wave periods LO-12 secs.). 


tere hl 


tlt MISTRESS 


ein 5 


me 


Fig. |. Map showing 


i cape Spences 


PATRICIA KOTT 


Water movement resulting from this swell is 
strong and pulsatile on the surface but 
decays with depth so that surge is moderate 
at 15 m and slight at 25 m depth. 

(2) “Sheltered Coast Subformation” (S.C.8.) 
(see Womersley & Edmonds 1958) refers to 
sheltered coasts where there is no swell and 
the coast is subject to waves of short period 
(up to 5 seconds) which decay rapidly with 
depth. Much of the coast-line in both 
Spencer Gulf and St. Vincent Gulf is of this 
type. 

(3) “Offshore Benthic” locations are those 
away from the shore where water movement 
results from tidal current rather than wave 
action. In St. Vincent Gulf tidal currents 
are generally about | m/sec., except over 
Tapley Shoal where they ate 1-2 m/sec. 


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locations in and adjacent to Spencer Gulf and St. Vincent Gulf, 


ASCIDIANS OF SOUTH AUSTRALIA 


These different locations. provide environmental 
conditions favouring an ascidian fauna of very 
varied habit. For each species its shape, or 
size. or mode of fixation, or growth characteris- 
tics appear to operate as selective mechanisms 
contributing to its success in the environmental 
conditions operating: 


(1) Jn Rough Coast Subformations, at depths 
less thin 15 m where surge is moderate to 
strong (Wright b, West Li, the dominant 
ascidian fauna is adapted ro the conditions by 
virtue of their colonial form, their vivi- 
parous larvae, their usually well-developed 
¢loaval systeros, and cither 
(ad un encrusting habit providing a large 
surface area for fixation (Atapozoa fanpea- 
siene, Cyxtodytes deHechiajel, Dideninum 
crendidum, Leptoctinides rufus, Lixsoclinisanr 
sf.  Oeulinaria australis. — Barrylleides 
nigrwen); ar 
(G} small stalks or sessile habit and cylin- 
drical body form enabling them to occupy 
sheltered crevices (e.g. on Frklonig hold- 
fasts. under Jedges, etc.). The stalks of these 
species are thick, and the colonics da not 
move freely with the currents (Podectevella 
cylindrica, Pseudodistomia cereum, Rilterelly 
herdimania, Syroicium papiliifertum) . 


(2) In Rough Coast Subformations, at 15 m 
und greater depths (Weight 1. West T.}, the 
strong surge at the surface ts reduced to mode- 
rile te slight water movement. Species with 
pliable stalks form a daminant component of 
the 4seidian fauna and are best able to exploit 
the changing direction of the water movements 
by moving with the water so that their bran- 
chial openings ate presented to the oncoming 
eurrent which thus reinforces the ciliary feed- 
ing mechanism. Only some ot these species 
have viyipurous lurvae (Borrylloides magni- 
coecus, B. leach, Palyearpa pedunculata, P. 
vlavata, Pyuru auytralis), 


(3) In Rough Coast Subformations, ut all 
depths, are large species fixed by a relatively 
small part of their surface. At shallow depths, 
they appear to be more offen on vertical rock 
fuces or in caves, where firm fixation can be 
uchieved, while at greater depths they are on 
the bottom (Polyciror giganteunt, Stvela pedata, 
Cremidecarpa erherldei, Herdntania mamus). 


(4) In Offshore Benthic locations with mode- 
rate currents and sandy bottums and some sedi- 
ment, there are: agnin, stalked species that are 
raised above che substrate und sometimes, bv 


we 


virtue of a pliable stalk. move with the current 
so that the branchial aperture is presented to 
the oncoming flow (A plidiuim colellnides, Pely- 
carpe clavate, Pyura sceresbiensis, P. spinifera. 
P, australis), 


(5) In Offshore Benthic Iecations with sluggish 

1o slow currents, dhere are: 
(a) Large species lying on of partly em- 
hedded in, of fixed to, rocky hottoms or to 
solid objects in sandy, often mobile. boi 
toms, These individuals and colonies are 
oriented. to take maximum advantage of the 
prevailing current flow by differential growth 
nf the colony or of the test, especially in the 
region of the siphons (Syeozan rerebyi- 
Jarmis, Aseidia spp.. Phallusia lepreysins- 
eala, Ctenieella  antipoda, Herriman 
miomus), Sycozea cerehrifarmis, which is 
wbundant on the bottom, especially in upper 
St. Vincent Gulf, has its “fans oriented to 
receive maximum current" (S. Shepherd, 
pers. comin). Its stalk is thick, short. and 
not pliable, and the species adapts to the 
direction of prevailing current flow by 
growth of the colony, In large sessile anil 
partly embedded species, the orientation of 
the siphons in relation to the current is 
effected by their Uifferential growth e.g, 
Phallusia depressivseulal, <Aseidia aclara, 
which is recorded only from sandy substrates 
in Which it is probably partly embedded, is 
especially interesting in the presence of cylin- 
drical tubes round the apertures cresting a 
Constimt Micro-environment. 
{b) Species with a leathery tesL sometimes 
produced into roots, in a sometimes mobil¢ 
sandy bottom or attached to the fibrous routs 
of the sea-grass Posidonia australis, These 
species often form aggregates of individuals. 
(Polycarpa pedunculata, Pyura irregularis, 
P. vinta, Halocynthia hispida, Micracosmrey 
Spp..).. 


(6) In Shehered Coast Subformations with 
shght wave action at the surface and Ho sedi- 
men, collections have been made from 3 to 25 
m. The species present represent all the groups 
previously distinguished: 
(4) Stalked species common jn Offshore 
Benthic locations and in Rough Coast Sub- 
formitions where there is moderate to slight 
surge, 
(b) Leathery aggregated specimens common 
at Offshore Benthic locations where the cur- 
tents are slight to sluggish. 


4 PATRICIA KOTT 


(¢) Aplousobranch species which, im more 
exposed conditions, are present in sheltered 
niches or crevices or have an encrusling habit 
(Podeclavella cylindrica, Distapla virlelis, 
Lepioeclinides rufus, Palvsyncraten orbi- 
ealren, Kehinoclinum verrilli, Ritterelle herd- 
miata, Svaoleume papilliferun), 


{d) The large stolidohranch and phleho- 
hranch species which exploit clean (vertical) 
rocky Substrates or protected tovations at 
Rough Coust Subformations and which are 
also’ present in Offshore Benthic locations 
where ihe current is slight. These large indi- 
vidupls are more often found at shallower 
depths and in less protected niches in these 
Shelicred Coast Subfermations than in 
Rough Coast Subformations (Ascidia spp. 
Rhedayema tarcievet, Corelle eumyeta, 
Herdmana momar). 


The presence of some of the larger phlebo- 
branch and stolidobranch individuals at shal- 
lower depths in certain areas where surge is 
greater, bul where clean stony substrate 1s 
available for settlement, suggests that it is the 
strengih of the current flow in telation ta the 
type of fixation which can he achieved that is 
the ¢ritical factor in site selection for these 
species rather than depth or light conditions. 
On the other hand, aplousobranch and stolide- 
branch encrusting species, and others whose 
shipe enables them to exploit narrow crevices, 
caves and ledges, appear to be affected more 
by fight and their depth range is more limited. 
These species accur at shallow depths both tn 
turbulent locations and in Sheltered Coast Sub- 
formations, and are not often taken in Offshore 
Benthic locations. They all have viviparous 
larvae and light sensitive organs which in- 
Ruence their settlement, and elficient achesive 
apparatus which ix needed where surge and 
turbulence is great, They are ulxo: common 
in ureas of gentle water Movement, tovether 
with the barge phlebohranch species not usually 
found at shallow depths in more Jurbulent 
areas, 


Seventy-six species are oow recorded from 
St. Vincent Gull and Spencer Ciulf. ‘This indi- 
cates a great diversity of ascidian species and 
suggests that conditions may be especially 
favourable for them. Records are mure nume- 
rous, however, from Gulf regions than from 
the “open” coast, probably because mure col- 
lecting has been done in these locations, It is 
not possible, therefore, with fhe infirmuation 
available, mm compare the faunal diversity on 


the open coast with that in Spencer Gulf and 
St. Vincent Gulf. 


Suborder APLOUSOBRANCHIA 
Family CLAVELINIDAE 
Subfamily Clave Lininac 


Clavelina baudinensis Kotr. 1957o0 S7_ Millar, 
146A: 3h3. 
New Records: Curickalinga Head, Rapid 
Hend. Previous Records: W Aust, (Rott- 
nest Island)—Kote 19574. Vico (Balnurring 
Beach, Laverton Bay, Williamstown) —Kott 


1957a; Millar 1966. Recorded from the 
intertidal ta G m. 


Deseripiien: Pwo ar more llal-topped lobes ol 
variuble size, joined by g common hase that 
is equal in height to that of the Iohes. Height 
of the colony to 4. cm, maximum diameter of a 
lobe 0.6 cm. The test is firm, getatinous snd 
transparent. Zoutds are blue. Thorax rounded, 
1.5 mm long; abdomen 7.5 mm long. with a 
well-developed pusterior abdominal  stolan, 
Zooids are parallel to the height of the colony. 
The branchial aperture, from the antetu-ventral 
corner of the thorax, is directed to the side. 
The atrial aperture front the antero-dorsal cor- 
ner of the thorax is directed vertically. There 
are 17 longitudinal muscles on each side of the 
body radiating from the apertures, 6 ventral to 
the branchial siphon, 7 extending along it, and 
4 extending along the atrial siphon, Dark pig- 
MEN spOls are present, anterior to, posterior to. 
and on either side of the base of the atrial 
siphon, ‘Fhere wre about 16 rows of about 30 
stigmata im the branchial sac. Nine obscure 
indentations are present around the margin of 
the branchial siphon, although the border of 
the atrial siphon is smooth and entire. The 
transverse vessels of the branchial suc expand 
into triangular languets as they cross the dorsal 
line. “The oesophugus is Jong. the stomach two- 
thirds of the distance down the abdomen is 
rectangular with 4 folds. Each zovid projects 
slightly above the Wat top of each colony. 
Gonads are present in the gut loop 


Remarks; Clavelina arafurensiy Tokioka. from 
the Arafura Sea. has similar colonies with 
zooids opening on the upper surface of the 
lobes, but is distinguished by the presence of 
distinct trunsverse nyuscles. Osyeoryriia fas- 
cieularis Tekioka, 1952, also has similar z00ids 
but there is x smueth stomach und zooids open 
all around a stalked head, thus’ distinguishing 
it from the present species. Two different types 
of larvae huve been described From specimens 


ASCIDIANS OF SOUTH AUSTRALIA 5 


previously ascribed (o this species, and it has 
been suggested (Kott 1969) that some colonics 
may in fact have been colonies of species he- 
longing to the genus Pyenoclavellu, distin- 
guished from Clavelina by the fertilisation of 
cgus al the buse of the oviduct. Those colonies 
with lurge numbers of evys at the same stage of 
development in the peri-branchial cavity and 
apparently fertilised there, belong to the venus 

Claveling as described, No other distinguish- 

ing character has been identified und us neither 

developitig eggs nor larvae were present in 
these colonies, this point has not been clarified, 

In St. Vincent Gulf the species is taken from 
sheltered locations where surge and wave uction 
is slight. The record from Rottnest t. (Katt 

19572) is From the intertidal area where it 

could sometimes be subjected to surge and 

wave action typical of the Rough Coast Sub- 
formation. [n such localities it would be found 
in shellered caves and crevices as it forms large 
soft colonies and is unlikely to occur in areas 
whee if is exposed {o sand or wave action. 

The red colour of the preserved specimen from 

Rapid Head is: probably the result of con- 

tamination from a sponge on which the speci- 

men was growing, as all other colonies ate 
bluish tn preservative, 

Podoclavella cylindrica (Quoy & Gaimard), 
Kot), 19574: 91. Millar, 1960; 64; 1963: 
716; 1966: 364. 
hada cylindrica Quoy & Gaimard, 1834; 


Claveline cylindrica. Michaelsen, 1930: 475 
and synonymy. 
New Records: West Beach, Hallett Cove, 
Port Noarlunga, Aldinga, West 1. (Oedipus 
Point), Wright [. Previews Records: W, 
Aust. (Albany to Rottnest f.1—Michaetsen 
$930; Kott 19574; Millar 1963. Vic, (Wes- 
termporl, Port Phillip Bay. Bass Strait}— 
Quey & Giaimard 1834: Millar 1960, 1943, 
1966; MacDonald 1858, 
FIG. 2 
Deseription; Zoids separate, joined by com- 
mon basal test into which postetior abdominal 
stolons extend. Occastonally zooids branch off 
around a central common axis (Wright 1). 
In immature colonies from Aldinga reef “drop 
off” there isa central vascular stolon extending 
up into each lobe and very numerous enlarged 
terminal ampullac surrounding the central ves- 
scl along its length, The abdomen may be 
equal to or less than the length of the thorax. 
When the thorax is contracted along the dorsal 
line, the oesophagus originates from half way 
along the length of the thorax. 


There es a dorsal pigment spot at the buse 
of the atrial siphon, and some pigment on 
either side of the dorsal line at the base of the 
branchial siphon, The atrial aperture is. ter- 
minal with a funnel-shaped siphon. ‘The bran- 


chial aperture extends laterally from the antero- 


ventral corner of the thorax. Ahaut 20 
Muscles cross the thorax obliquely front the 
ventral to dhe postero-dorsal cormer of the 
thorax anc continue along both sides of the 
abdamen. When the dorsal line of the zoids 
is strongly contracted, the muscles an the \horax 
lie almost at right .ngles to the rows of stig 
mata. The oesphagus is long und there is 4 
presiomach swelling halfway alone {ts length, 
The stomach is large and square, Clumps of 
18 or more embryos ure present in brood 
pouches formed at the postero-dorsal corner of 
the thurax. Gonads are present in the gut loop, 
Larvee: About 1.2 mn lang. Anteriorly there 
is at Hat frontal plate bearing three adhesive 
papillac with uccessory cup. arranged ino 
triangle. The larval thorax is characteristically 
deep. 

Remarks: This species is especially common. 
The relatively short.ahdomea, the prestemach, 
the form of the colonies, and the presence of 
pigment spots on the anterior part of the thorax 
are characteristic, 

The colonies flourish only in protected caves 
or crevices and venerally from vertical faces in 
sreas where there is no sill or sediment. In the 
Rough Coust Subtormation, the species is 
found at depths of 10-22 m, and in the Shel- 
tered Coast Subformation at 3-10 m deep, 


Podoclavella moluccensis Sluiter, 1904; 5, 
Hastings. 1931: 82 and synonymy. Katt. 
1963: 90. 


New Record; Tipara Reef (Spencer Gulf). 

Previous Records: W, Aust, (Cape Boileau. 

Garden Island, Rottnest 1.)—Sluiter 1895: 

Kott 1963. S Aust. (Port LincoIn}—Kott 

1963. Qld. (Great Barrier Reef}—Hustings 

1931. 

FIG. 3 

Description: The colonies form extensive mats 
consisting of a basal membrane sUpporting 
dense array of upright lobes, each consisting of 
a single zooid enclosed in a soft transpyrerit 
test. Occasionally the basal half of adjacent 
lobes is Fused, The zooids are pale to dark 
bluc and there ts no specinl accumulation into 
specific pigment spots around the apertures. 
The zooids are cinsely acherent fo the test 
und extend the full length of the free lobe for 


6 PATRICIA KOTT 


fen" 
SA ese e 
aay Nes 2: 


Fig. 2. Podoclayella cylindrica, (Hallett Cove, 8 m). Colony. 

Fig. 3. Padoclavella moluccensiy. (Tipara Reef), Thorax showing muscles. 

Figs. 4 Atapozoa fantasiana, (Wright I.). Fig, 4.—Contracted zooid. Fig. 5—Zooid with brood 
pouch and embryo. 

Figs. 6, 7. Distaplia viridis. (Reef off Hallett Cove, 8 m). Fig. 6—Zooid with mature 2 gonads and 
brood pouch, Fig. 7.—Zooid with mature ¢° gonads. ; 

Figs. 8.9. Polycitar givanteurn. (Port Noarhinga), Fig, 8—Immature larva. Pig. 9.—Mature Jarvis. 

Figs. 10-12. Endistoma renieri. (Wright T.. 10 m), Fig. 10—Zooid, Fig. 11.—IJmmature larva. Fig. 
12.—Mature Jarva. 


Ww 


ASCIDIANS OF SOUTH AUSTRALTA 7 


their whole length, The atrial wperiure is ter 
minal and the branchial aperture from the 
anteru-ventral part of the thorax is inclined ar 
a slight angle to it but is not rectirved, There 
are ybout 30 transverse muscles extending from 
the ventral to the dorsal border of the thorax 
and anaslomosing with one another both ven- 
irally and dorsally. About 6 of the mast 
antenor Iftunsverse muscles extend from the 
short siphons ta cross the dorsal line. The 
most posterior transverse muscles terminate 
around the region of the oesophagus. No 
muscles were detected on the abdomen, There 
ate 17 rews of ahowt 50 stigmata. There is a 
small prestomach enlargement half way down 
the oOcsophagus. Whe stomach is smooth 
walled, large and rounded half way down {he 
abdomen, 


Remarks: The specimens are easily confused 
with Podeclivella cylindrica, fram which P. 
moluccensis ts distinguished by the extensive 
basal. membrane, the absence of o recurved 
branchial siphon, the very large number of 
transverse muscles which do not extend along 
the abdomen, by the close adherence of the 
body wall to the test. and by the absence of dis- 
tincl pigment spots around the apertures. 
Shepherd [pers. comm.) states that this 
species at Tipura Reef is seasonal, appearing in 
early winter aid dying off during carly summer. 


Subfamily totozoiae 
Ataporoa Fantasiana (Kort) 
Biassrsrai fantasiana Rott, 195%a; 76; 1967; 
1A? 


New Record: Wright I Previous Records: 
S. Aust. (Reevesby [.)—Kott 1957a. 
FIGS. 4, 5 

Pescription; Flat irregular investing colonies 
about 03 cm thick. Test soft, jelly like, semi- 
transparent. Both apertures of zonids open 
separitely to the extenor. The postero-dorsal 
aspect of the peribranchial cavity is expanded 
into a brood pouch with two embryos at differ- 
ent stages of development. Black pigment is 
scattered throughout the test, but the colony is 
a light purplish colour. Zooids up to 3 mm 
in length. Zooids have 16 to 20 fine Jongitu- 
dinal muscle bands forming a wide open mesh- 
work with the transverse bands on the tharax, 
There are 3 rows of up te 25 clongate stigmata; 
the ocsophagus is long, the stamach smouth 
and oval, and there is a rounded posterior 
stomach. The apertures ate stall end the 6 
lubes of the margins indistinct 
Lurvee; Latge. as previously described, with 


characteristically clomgule ureas of adhesive 
cells. 

Remorks Even in the absence of the com- 
pletely distinctive larvue and brood pouches, 
the species is characterised by the closely set 
apertures and short alrial siphon, by the open 
meshwork of muscles on the thorax, by the 
comparatively short zooid, and by the very 
large number of stigmata in euch row. 
Distaplia viridis Kott, 19570: 96, Millar, 1966: 

365, 


New Recordy: Hallett Cove, Port Noarlunga 
Reef, Carickalinga Head. Previous Record 
S. Aust. (Victor Harbour, Reevesby I.) — 
Kot '957a. Vic. (Port Phillip Bav)— 
Millar 1966. 


FIGS. 6, 7 


Description: Living colonies from Hallett Cove 
had a transparent matrix with orange zovids, 
While these specimens ate greenish in preserva- 
Uve due to the greenish colour of the enclosed 
zooids, Preserved colonies from Port Nonr- 
lunga are also greenish but the living colonies 
were blue-black with white markings, ‘Test is 
semi-transparent and very soft, Zooids closely 
pliced more or less in double rows. Colonies 
are izregular and investing, about 4 mm thick. 
The surface is always smooth. There are no 
sund inclusions. Common clouedl upertures 
are randomly distributed over the sutface of the 
colany and zooids are arranged on either side 
of very shallow and narrow common cloacal 
canals, A brood powch is developed from the 
postero-dorsal corner of the thorax and con- 
tuins only a single embryo. ‘The atrial lip is 
sometimes tridentate at the tip with a longer 
median lobe, This, however, may be obscured 
if the atrial hp is widely extended. In younger 
colonics the zooids may be in circular systems 
of 5 to 14 zovids. There are 4 rows of stle- 
muta with, parsestigmatic vessels. The sin- 
mach has glandular folds internally but exter- 
nally is smooth. There are ahout |) testis lohes 
in-a rosette in the loop of the gut, and a single 
ceg protrudes from the right side of the ahdo- 
men, A Conspicuous gastric reservoir is also 
present in the loop of the gut. 


The single embryo present in the hrood 
pouch jy as previously described. The tail of 
the larval form is especially short and extends 
only half way along the ventral surface, The 
larval test has a foamy appearance. 
Remarks: The species conforms with speci- 


mens previously taken from Victor Harbour 
and Reevesby I., South Australia and. the pre- 


¢ PATRIA KOTT 


served colonics have the same erecnish tinge 
in formalin resulting from the colour of the: 
zooids. Colours present in the living speci 
mens, however, appear to vary The single 
einbryo in the brood pouch 1s apparently 
characteristic of the species which is common 
in St. Vincent Gulf and Spencer Gull although 
\t has nat heen recurded from other localities. 


Sycozoa cerebriformis (Quoy & Guaimard), 
Brewin, 1953; 58 and synonymy. Kott, 
1957a: 99. Millar, 1966: 365, 
dutialie verelrifermne Quoy & Gaimard. | 834; 

New Rererds: Off Troubridge 1. Orontes 
Bank (off Port Vincent), upper St, Vincent 
Qiull, Hallete Cove, Canckulinga Head. West 
1 (Toad Head), Wright 1. Previews Recordy: 
North-west Aust.—Hartmeyer 1919, 8. Aust. 
«Victor Harbour, Port Lincolnj—Koatr 
57a, Caullery 1908, Vie. (Balnarnng 
Beach, Westernport, Point Lonsdale )—Quoy 
& Gaimared 1434; Caullery 1908: Michaelsen 
1924; Kott 19579; Millar 1966. N,S.W. 
(Gunnamatta Bay, Jervis Bay, Port Jackson, 
Port Stephens} —Herdman 1899; Kolt 
t9S7u. South Africa—Hartmeyer 1912; 
Michaelsen 19230 


Description: Colonies {rom fun-shuped 10 
curved lamellac. Zooids arranged in double 
rows down both sides of these lumellac, bran- 
chiol apertures opening to the exterior 
Cloacal apertures, however, us is usual in this 
genus, open into common cloacal canals ex- 
tending Vertically down both sides of the 
colony. These cloacal canals open separately 
wround the edge of the marrow fat top of the 
colony, 


Remarks; In Sycozed sigillinoides Lesson, from 
the Aataretic (sce Millar 1960; Kott 1969), it 
has been found that the cusmmon cloweal cuvi- 
ucs open into a ring canal round the anterior 
end of the cofony und this ring canal is part 
ef a common cloacal cavity opening by a ter- 
minal aperture. Brewin (1953) characterised 
the genus Sycozed by the condition of the 
cloacal cunals opening separately around the 
anterior border of the colony, Both Millar 
(1960) and Kott (1969), working with speci- 
mens of Svcecna sigillineides from the Ant- 
arctic, did not accept this interpretation of the 
cloacal openings and suggested that Brewin's 
caloni¢ts were distended to expose the openings 
in the cloacal cavity. Brewin's observations 
for both Sycoezea cerefrifernis and far §. 
tenuicaulis are accurate. The situation in &. 


sigillinoides, however, tndicates. Lhat sepurate 
openings of the canals is pot a character shared 
by all species of the venus Sycozeu, 

Colonies have been observed with their wide 
fins, from the short, sturdy stalk, oriented to- 
ward the oncoming current (S. A. Shepherd 
pers. comm.). ‘The stalk is nor Meaxible. as in 
S, renaicaulis, and the orientation of the colony 
is unlikely to ulapt 10 changes in direction of 
current flow. The species ts most cummon 
attuched to shell or rock surfaces at locations 
where there are slow to sluggish currents, and 
where the light intensity is not great due to 
depth and sediments, Larvae have an otolith 
bit no ocellus (Caullery 1908), They have 
relatively short tails, their tree swimming exis- 
tence is short and they are probably not strong 
Swimmers. 

The species therefore is well adapted to an 
existence in lucytiony with slow ta sluggish 
currents where it 3s most commonly found, The 
low fight intensity al these stations, due to 
depth or sedimem, is coincidental and not 
likely to directly uffeet settlement of these light- 
insensitive Jarvac, 


The species is taken from the Rough Coast 
Subformation at West |, and elsewhere in con- 
ditions of moderste surge, either at depth or in 
crevices, or under houlders where it ts pro- 
lecled, Again. the low light intensity is only 
coincidental with The occasional occurrence of 
this species in these situations where light sen- 
sitive aplousohranch larvae that are attracted 
mito shade are more common. 


Sycozoa tenuicaulis (Herdman), Brewin, 1953: 
57. Rott, 195%a: 99. Millar, 1963: 707. 
Calella tennivantis Herdman. 899, 64, 

New Record: Off Broadway. Previews 
Rereords; W. Aust.—-Millar 1963. Vie. (Port 
Phillip Bay. Lakes Entrance!—Koll 1957. 
Millar 1963. ‘Tas. (D°Entrecasicux Channel, 
Furneaux = Group)—Millir 1963: Kort 
1¥57a. NS.W. ¢ Botany Bay, Jervis Bay, 
Broken Bay, Port Stephens, Part Jackson) — 
Herdman 1899: Millar 1963; Kott 1957a, 


Descriprian: A single colany only is available 
und is the usuul flattened inverted cone. fixed 
by a long stalk with busal hair-like rootlets. 
Zooids are present in closely set double rows 
along the length of the head. ‘The longitudinal 
common cloucal canals extend the length of 
the head between each double row of zooids 
and open by a wide opening around the out- 
side margin of the flat top of the head as pre- 
viously described by Brewin (1953), 


ASCIDIANS OF SOUTH AUSTRALIA 9 


Remarks: The observations by Brewin on the 
separate cloacal openings around the top of the 
head are confirmed in the present colony. The 
species is distinguished from the superficially 
similar Antarctic species. S_  sigillineides 
Lesson, by these separate openings of the com- 
mon cloacal canals, which, in S, sigillinoides 
open into a terminal chamber with a single 
common cloacal opening on the centre of the 
upper free surface of the head (Millar 1960; 
Kote 1969). The species are also distinguished 
by the flattened head and by the tuft of hair- 
like roots in §. tenuicaulis (see Millar 1963). 


Records of §. tenuicaulis are confined to 
Australia, and at present the species is known 
only from fairly protected bays. It is possible 
therefore that its isolution has resulted in spe- 
ciation separating it from the more widely dis- 
Uibuted circum-polar 8. sigi/linoides. The fat- 
ter is also known from South Australian loca- 
lilies (Kot 1969), 


Brewin (1953) states that all records of this 
specics are from deep water, Although this is 
not strictly accurate, there are indeed no 
records available from the inter-tidal region. 
Specimens. have. been taken from a depth of 
4m (Millar 1963) ta 50 m (Kott 1967}. 


Shepherd (pers. comm.) has observed that it 
is fairly common at sub-littoral locations in 
deeper water, with tidal currents up to 0.5 
m/sec, fone knot). The larva of this species 
does not have @ light sensitive occllus (Brewin 
1953) and would he at a disadvantage in seek- 
ing suitably protected locations for settlement 
in waters where there is apprectable wave 
uction or surge but, like 3. sigillineides (see 
Kott 1969), ts well adapted for an existence on 
the sea floor. 


Family POLYCITORIDAE 
Polycitor giganteum (Herdman). 


Polyclinum giganteum Herdman, (889: 79. 

Polyclinuni globosum Herdman, 1899: 80. 

Polyoltor gelatinnsa Kort, 1957a: $3. 

won Palycitor gigantewn Slviter, 1919: 10 

{Diazona siganteum Sluiter). 
New Records: Tapley Shoal, Hallett Cove, 
Pon Noarlungs, Aldinga, West I, (Toad 
Head), Wright I, Previous Records: W 
Aust, (Rottnest 1). S. Aust, (Port Noar- 
junga). Vic. (Balnarring Beach, Lakes 
Entrance, North BrightonJ—Kott 1957a. 
N.S.W. (Jervis Bay, Port Tackson}—Herd- 
man 1899; Kott 1957s. 


FIGS. 4, 9 


Description; Large. fan-shaped er rounded 
lobes of varying size; sometimes smaller lobes 
occur together fixed to a common base, The 
test is firm, gelatinous without sund inclusions. 
and is semi-transparent and almost glassy in 
appearance. Zooids cun be seen radiuting from 
the basal constriction of the colony to oper an 
the rounded upper surface. Living zooids are 
cream to bright orange but are pinkish in pre- 
servalive. The diameter of the colony js gra- 
dually reduced toward the base where it is 
fixed to the substrate. In the colony from Tup- 
ley Shoal, two lobes branch from a comtnon 
base and the test of the upper purt of exch inbe 
is coalesced. There are [5 longitudinal 
muscles pet side continuing as 3 bands along 
each side of the abdomen. The stomach has 
4 folds and there are 10-!2 rows of 22 to 40 
stigmata, 


There are 3 to 9 developing embryos in the 
atrial cavity and in the distal portion of the 
oviduct. Larvae are large, about 1.2 mm, anil 
large ampullae develop around the base of the 
3 median papillae as previously described for 
&. giganreum, 


Remarks: A teexamination of the type speci- 
méns of P. gelatinosa from Rottnest 1.. has 
shown that che colonies are slightly smaller 
thatt most colonies of FP, giganieum. The zooids 
and the test are, however, identical with those 
of P. givanvevem. Further investigation of 
larvae from typical colonies of P, giganteum 
has also shown that in the less mature Jarvae 
the anterior ampullae are not developed and 
these Jarvae appear identical with those des 
erthed for P. geluinosa (Kott 19$7a)l. As 
there is so much variation in the shape and size 
of colonies of P. pigamteunt. from spherical 
individual lobes to numerous pyriform lobes 
from a common base, this cannot be regarded 
as a valid character on which to separste the 
two species. The gelatinous test, large zooids 
and larvae are characteristic. 


Fudistoma pyriforme (Herdman), Hastings, 


1931; 84. Kort, 1987a; 75, Tokioka, 
1950: 120; 1967: [1O. Vasseur, [969- 
918, 


som maplidtures pyriforme Herdman, (886: 
19. 


New Record: Off West Beach. Previous 
Records; S. Aust. {Port Noarlunga)—Kott 
1957a. Old. (Great Barrier Reef, Flinders 
Passage)—Herdman 1886. Pacific (Palaa 
Is,, Gilbert 1,)—Tokioka 1950, 1967. Indian 
Ocean (Madagascar)—Vasseur 1999, 


1h PATRICIA KOTT 


Description; Rounded lohes. narrowing to- 
watds the base where the test expynils into a 
basal plite From which several heads may rise, 
Sund ts ubsent from the outer 5 mm of test 
on the upper half of the colony but is present 
internally and is also present through the test 
in the basal half of the lobes, Maximum dia- 
meter of head is 4 cm, The test is firm and 
gelatinous. The colony is grey in preservalive, 
Zooids are present, upening over the upper sur- 
face of the head. Thev are arranged in circu- 
lar systems, with the alnal apertures opening 
separately in ua cirele in the centre of the outer 
circle formed by the branchial openings. Each 
atnal aperture is protected by a lobe of test 
that covers (he opening from its dorsal surl'ace 
and it appears that the excurrent stream from 
each yooid would he directed towards the 
centre to reinforce the excurtent stream from 
zooids in the saine circular system, The incur- 
rent ciliary stream is probably drawn from an 
grea immediately adjacent to the branchial 
aperture, This arrangement of apertures tep- 
resents a Stage before the development of truc 
cloacal systems. 

Zonds are 5 10 7 mm long, of which the 
thorax is only | mm, They cross one another 
in the test. Both siphons are well developed, 
anteriorly directed, and are surrounded with 
circular muscles to form a distinct sphincter. 
The atrial sphineler i especially well deve- 
Inped, There are about 20 longitudinal muscles 
on the thorax. although these may be reduced 
te LZ in contracted specumens. The trans- 
verse musculature ts fairly strong. 

There wre 3 tows of about 9 to 12 stigniti. 
he stomach is smooth and tounded and in 
contracted specimens the intestine behind the 
stomach forms jin “S" bend asx previously des- 
uribed for this species. The rectum forms the 
ascending limh of the gut loop, 


Remarks: Specimens of Mudistoma are noo- 
riowsly difficult to characterise and the variable 
condition of the intestine in the present speci- 
mens suggests thal this feature, previously 
regarded as a diagnostic feature, is dependent 
on the degree of vontractian of the abdomen, 
Speciniens identified as EF, pyriforme from 
Heron band North West I. (Capricorn Group) 
have been examined. Zooids are arranged in 
similar systems to those described nhove, 
although these may be obscured hy sand in the 
surfuce lest; the proximal part of the intestine 
forms either an “S" hend er a loop. and pig- 
meni is presen is spberical cells in the surface 
lest. Despite the variation in the external 


appearances of these colonies they all appear to 
belong to £. pyriforemne, characterised mainly 
hy the condition of the thoracic miuseulature, 
the lofig oesophagus, the atrial sphincters and 
the arrangement of zooids in the colony. These 
characters arc. te some extent. shared by other 
specics and it is possible chat more than a single 
species ts represented hy the records ascribed. 
lo this specres. 


Enudistoma renieri (Hartmeyer). Michaelsen, 

1923: 10, Kom. W57as 74. Millar, 1962: 

160, 

Polveitar renier! Hartmeyer, '9)2: 39. 
New Record, Outsile Woght BE Previews 
Records: W, Aust. (Point Peron) Koti 
1957a. South Africa—Hartmeyer 1912: 
Michaelsen 1923a; Millar 1962. 


FIGS. 10-12 


Description: Fleshy investing colany, 0.6 ¢m 
thick. Test semictranspurent with reddish to 
black pigment cells in streaks on the surface. 
The surface of the test is smooth, without 
foreign bodies or sand, and is depressed over 
the zooids. Zooks are arranged in circles of 
about 4 mm diameter, the branchial openings 
around the periphery of the circle and the atrial 
openings toward the centre, protected by lobes 
of test. The atrial openings are in a pigment- 
free area, The zooids do not cross one another 
in the Lest. The abdomen is about twice the 
length of dhe thorax. “Che atrial aperture is on 
w evlindrical siphon which js about three times 
the length of the hranchial siphon. The body 
wall is fairly muscular with at least [2 longi- 
tudinal muscle hinds of 4 to 5 strands crossing 
numerous transverse bands. The longitudinal 
binds appear to separale our into separate 
strands. When not so strongly contracted, the 
ciraar muscles. around the atfial siphon are 
strong and conspicuous ulthough they ute 
sprend along the siphon rather than forming a 
large sphincter muscle. There are about 20) 
long rectangular stigmata in each row. The 
rounded smooth stomach is halfway down the 
abdomen, There is a long duodenal urew and a 
short round posterior stomach, The part of the 
intestine distal ta the stomach 1s sometimes 
kinked in contracted specimens. The gonads 
ure in the gut loop. There is an expansion 
from the dorsal uspect of the posterior end of 
the thorax accommodating a loop of the avi- 
duct with one to two embryos, and although 
the brood pouch is not separated from the 
thorax by a narrow stalk as in the true brood 
pouch of the Holozoinae, jt is structurally 
homologuts, 


ASCIDIANS OF SOUTH AUSTRALIA V1 


The larvae are about | mm long, typically 
polycitorid, with the 3 median papillae develap- 
iag on short. stalks fram <lepressions in the 
centre of rounded swellings around the anterior 
end of the larva. The margins of these depres- 
sions become attenuated in the mid-line to 
form median ampullae at the base of the papil- 
Jury stalk. The area of adhesive cells in these 
papillac is lengthened tongitudinally to different 
exlents lor each papilla, This lengthening is 
Teminiscent of the condition in Atapezaa 
larvae. 


Remarks: Distinctions beeween Enaisrama spp. 
afe not allogether sulisfuctory and many 
characters such as the body musculature, length 
of gut. and looping of the jntesline, all vary 
with the degree of contraction of the body. 
The present species is identified by the gela- 
linous nature of the fest, by the large number 
of stigmata, by the long oesophagus and the 
position of the stemnch mid-way down the 
abdomen, The extended adhesive urey of the 
larval papillae was not recognised previously 
(Kott 1957a). A re-examination af Kolt’s 
specimens from Point Peron, Western Austra- 
lia, has demonatrated that the papillae are iden- 
tical with those in the present collection. This 
character therefore appears to be distinctive for 
the species, 

The zoonis of the Australian specimens 
resemble Hartencyer's (1912) South African 
specimens, although the colony of the South 
Austialian specimens is thinner.  Millar’s 
(1962) specimens appear to differ in many 
chatacters, however; notably in the reduced 
size of the thorax, in the position of the sto- 
mach at the posterior end of the ahdamen, in 
the mumber of muscle bands and rows of stig- 
mata in the length of the atrial siphon, and in 
the cylindrical form of the colony. 


Cystodytes dellechiajei (Della Valle), Kott, 
1954: 154 and synonymy. Tokioka, 
1930: 120. Millar, 2953: 284; 19H0: 82; 


1962; 143; 1963: 713: (966: 34S. 
Distoma deNechiajiae, Della Valle. 1877: 40, 
? Aplidiam lobatium, Delle Chiaje, 184. 390 
(not Savigny 1816). 

Oystadytes dellachiaiae. Kott. 1987a: 68. 
Cystodyies Delle Chiajer Pérés, 194%: 171. 
New Record: West 1, (near Penguin Rack) 
Previous Records; W. Aust. (Dampier 
Archipelago to Alhanyi—Michaelsen 1930- 
Kott 1954, 1997u; Millar 1963. Vic. (Port 
Phillip Bay, Barwon Heads}—Millar 1946. 
Tas, {Maria T,.)—Katt 1954. Pacific (Palao 
Is.)Tokinka 1950. New Zealund (North 
L. Chatham  [s,}—Michaelsen 1924; 


Brewin L948, 1951, 19529, 1956; Millar 
1960, California (Coronado 1, Puerta 
Esconido!—-Van Name 1945. Indian Ocean 
(Ceylon)—Herdman 1906. Meditteranean 
—Della Valle 1877: von Drasche (883; 
Lahille 1890; Harant 1925, 1929. Afncea 
(Mozambique, Gold Const, Cameroons. Sene- 
gal)—Michaetsen 1915: Pérés 1948: Millar 
1953. 1962. The species is also known from 
the Atlantic Ocean, along the east coast of 
the American continent from Patagonia 
éMillar 1960) to the Caribbean and from the 
Azores (Michasisen £9230), the Canary 1 
(Hortmeyer 1912) and Virgin Is. (Van 
Name 1945). ft has been tuken intertidally 
and 10 uy maximum depth of 736 m (olf 
Brazil, Herdman 1886)- 


Description: Iwregular investing colonies, Liv- 
ing colonies purple with colourless "splotches" 
but in formalin the colonies are brown with 
white blotches where zooids are present |n the 
test surrounded by the calcareous spicules thil 
ure typical of thix specres, The species is espe- 
cially constant and the present colonies and 
zooids conform exactly with previously des- 
cribed spectmens. Larvat are present in brood 
pouches attached to the parent zooid or free 
in the test, The larvac have the usual large 
papillae surrounded by <ctodermal ampullie 
which have coalesced distally to form a circle 
around the papilla as described previously for 
the species (Kott 1954. 1957a). 


Tumily POLYCLINIDAE 
Subfamily HUHERPMANUNAE 
Ritterella herdmania Kott, 1957a; 102 (nom. 

hov.); 1963; 78 and synonymy. 
New Rererd: Port Noarlunga.  Previaus 
Records: W. Aust. (Green Pools)—Kott 
STi, NSW. (Newport, Port Jackson, 
Wattamolla}—Herdman 1899: Katt 1957a. 
1963, 


FIGS. 13-17 
Desctiplion: Sandy finger-like lobes joined 
basally, The lobes are long. and slender, spoon- 
shaped terminally, with 1 to 5 zooids in each 
lobe. The branchial apertures open into the 
cancavity of each lobe and the atrial apertures 
open round the convexity of the anterior tip 
of the Jobe. Both apertures are 6-lobed and 
on very short siphons, The branchial aperture 
is terminal and the atrial aperture rises from 
opposite the first row of stigmata. There are 
circular siphonal muscles, very delicate longi- 
tudinal muscles und some weak transvetse 


12 PATRICIA ROTT 


hiuseles on the thorax, There are five rows of 
h to 16 stigmata in the branchial sac, some- 
times. in the larger zooids, parastigmatic ves- 
sels re present in some of the rows of stizg- 
Mats and appear to brsect them horizontally 
to fori éxtra rows. Triangular langucts are 
present in the mid-dersal tine expamded Crop 
both the transverse vessels and the para-stig- 
matic vessels. Smaller rounded papillae are 
wis present in the middle of cach transverse 
yessel on either side of the branchial sac. These 
papillae have not previously been described 
for this venus. The fact that they do not arise 
on the parastigmatic vessels suggests that rhey 
may be present us rehets of pupillze supporting 
longitudinal vessels in the branchial sac anc 
homologous wath che papillae present in’ the 
Anturetic genus Tylobranchion. 


The condition of the stomach varies aecord- 
ing to its degree of contraction and when 
extended there ure apparently four to six 
stomach folds. but these are not always dis- 
tinct, Four folds sometimes appear to be pre- 
sent only in the anterior part of the stomuch 
There is iso a small posterior stomuch us pre- 
viously described, The posterior abdomen may 
be very Jung and thread-like ine testis follicles 
are arranged in it inva single row, The ex- 
tended thorax und abdomen together measure 
4-5 mmi. The posterior abdomen is consider- 
ably longer 


Larvie are present in the thoracic cavity of 
some of the zooids. They have 3 anterior 
papiilae in the median line alternating with 
paired anterior ampuilae. Dorsally and ven- 
trully paired rows of ampullury vesicles extend 
posteriorly. There is an otolith and acellus. 


Remarks: Uhe variations in the number of rows 
of stigmata resulting: [ram then’ bisection by 
purustigmatic vessels and the increase in the 
size of the zovid-byaring lobes, both of which 
oecul with increasing maturity, steests that 
confusion could arse regarding the identity of 
specimens assigned to this and to related. spe- 
gies, Part olf the type colony of Riverella 
wyoenterrica Millar, 1966, from Port Phillip 
Bay, has been examined, The external appear- 
ance of the colony resembles &. herdmenia and 
the 10 rows of stigmata could have resulted 
from the bisection of 5 primary tuws hy para- 
stigmalic vessels, as the triangular dorsal lan- 
gucts arc of two alternating sizes. There are 
no papillae on the transverse vessels in Millar's 
species, however, und the stomach folds are 
also distinctive. 


Vive primary rows of sligmata appear to be 
characteristic of most Rivere/la spp. although 
the number can be merensed probably by sub- 
division with parastigmatic vessels which sub- 
sequently are not distinguished from printuiry 
transverse vessels.  Rirrerella herdmanin, R 
eeduncilata Tokioka and Ro vesteta Millar, 
1960 (from North [., New Zealand) have para- 
stigmatic vessels and sometimes increased num- 
bers of tows of stigmata: R. proliferix (Oka) 
(>R. dispar Kott, 1957a) from Japan and 
from the central east coast of Austealia (soc 
Tokioka 1953a; Kott 1957a, 1963). and R. 
vigillinaides Brewin, 1958u. from Stewart T., 
have only the 5 primary rows of stigmata und 
noo parastignyatic yessely, Re asvenerrica 
Millar has increased numbers of rows of stig- 
Mata and apparently no parastigmatic vessels. 


The type species of the genus Exherdmania, 
E. claviformis (Ritter) (see Van Name 1945), 
together with &. selidy Millar, 1953 fram the 
Africeun Gold Coast, E. virrest Millar, 1961 
fram Brazil, and E. afgitarad Millar, 1963 from 
northwestern Australia are easily distinguished 
by a Jong aesophagus, a large number of rows 
of stigmata and the absence of parastigmatic 
vessels and, where ther larvae are known, by 
the modified adhesive organs as described for 
this genus and for Placenicla spp. (Rou 1969), 
Kitherdmunia australis Kott, 1957a, however. 
fram South Australia, Wietoria and New Sanath 
Wales, has a short ocsaphagus, 12 10 19 rows 
of stigmata, purastigmulic vessels, aimst is papilla 
in the middle of the transverse vessels on cuch 
side of the body. It as distinguished fram A&. 
herdmania by the single zooid in each lohe of 
the colony, the ahsxence of stomach folds. the 
number of rows of stigmata and the testis fel- 
licles which wie bunched in the posterior abdo- 
men. 


Larvae are known for R. prolifervs and R. 
herdmania, and are typically palyclinid with 
ampullary vesicles. 

Tn the present species and in K. australiy the 
papillae on the transverse vessels ate remini- 
seent of Tylobranchion and felated genera, 
and probably represent a primitive character. 


Pseudodistoma cercum Michaelsen, 1924: 364. 
Kou, 1963: 77 and synonymy. Monniot, 
1969; 437 

New Record: Nora Creina Bay. Previous 
Records: NSW, (near Eden) —Kote 1963. 
New Zealand (Stewart I. (Paterson Inlet), 
Foveaux Strait, Otsgu coast, Lillle Papanui, 
Great Barrier 1)—Michacisen 1924: Brewin 


ASCIDIANS OF SOUTH AUSTRALIA i3 


1950e, MY58a. Atluntic Ocean ( Dakar}— 

Munniot 1969. The species t& known inter- 

tidully and down to 87 om, 

FIGS, 1k, 19 

DPesiprian: Soft. gelulinous, semitesnsparenr, 
rounded or evlindrical hens of slightly preater 
diumeter than the more leathery stalk of up to 
5 cm leneth. In some specimens the stalk is 
expanded into yu thick mut from which 
humerous heads arise. The zooids are 
numerous and cpen all around the head by 
separate G-lobed branchidl and atrial openings, 
The contracted thoray and abdomen together 
measure only 2 om. Fine jongitudinal muscle 
bunds on the thorax mumber 20 to 30 and 
these extend along both sides of the abdomen. 
There are 15 10 20 pows.of stigmata in each nf 
the 3 rows, The 4 stomach folds are obscure 
and may be artefacts resulting from the col- 
lapse of the stomach. A duodenal awelling and 
wu rounded posterior stomach are also present. 
There is a long ovary, with numerous eges 
more than halfway down the abdomen, but no 
temis follicles were present in the colonies from 
these stations, There is a single developing 
embryo in a brood pouch from the postero- 
dorsal corner of the thorax. 


Renurks; The generul form of the colonies. 
arrangement of body musculsture, the bran- 
chial sac, gut and the situation of the ovary 
some distance down the nosterior abdomen, all 
agrze with the previously described specimens. 
All other species of the genus have a similar 
situation for the ovary some distance along the 
pesterior abdomen: P. efricatigm Millar. 1954, 
1%62, P. fragiliv Tokioka, L9SR; P. evrnetsense 
Péres, 1952: P. antinboia Tokioka, 1949; S. 
wpaca Brewin, 1950¢, P. Arieni Péres, 1949. 
The stalked colonies of P. africanum are also 
reminiscent of the present species in the pre- 
sence of a single developing embryo in a 
thoracic breed pouch and ace distinguished 
only by a smuller number of longitudinal 
thoracic muscles, As there has been consider- 
able variation demonstrated in this character, 
the distinction is suther doubtful, and the spe- 
cies Of its telatives appear to have a wide cir- 
cumpalar distribution in the southern temperate 
region as Monniot (1969) has ulready inei- 
cated, 
Subfamily PocycuNtwac 
Aplidium pliciferum (Redikorzev) Kort, 1963; 
106 


Amaroticii pliciferim Redikorzéy, 1927: 
39D, Tokioky, [953a; 183; 1962: 2: 1947; 32, 
Aplidine phortax, Millar, 1966: 359. 


New Records: Troubridge Shoal, Hallett 
Cove. Previous Records: W. Aust. (Point 
Peron, Rottnest 1.)—Kout 1963. Vio. (Port 
Phillip Bay)—Millar 1966, Japan (coastal 
water of Honshu, Shikoku and Kyushi and 
the Inland Sea)---Redikorzev 1927; Tokioka 
1953a, Hawaiian Is. (Auau Chunnel)- - 
Tokioka 1967. 
FIG. 20 
Description: Rounded, soft, sessile colonies. 2 
cm in diameter. Iq life the colonies are bright 
yellow. The surface of the colony has deep 
Jurrows marking it off into extensive rounded 
areas with up te 2 common cloacal openings 
from which double row systems muiate Test 
transparent. zooids orange in the living speci- 
men. Thorax and abdomen wre of cyual length 
and together meusure 2.5 mm. The posterior 
wbdomen is long, up to 8 mm. There sre 4 
well-defined branchiai lobes. a strong circular 
branchial sphincter and & fine longitudinal 
muscle bunds which extend down cuch side of 
the thorax, The upper border of the atrial 
opening is extended inta a small pointed lip 
sometimes tridentate There are 8-10 rows of 
about 19 stigmata, The oesophugus is long and 
the stomach, about half way dawn the abde- 
men, has 19 to 25 well defined folds. There is 
w duodenal swelling and a small posterior 
somuch, Two developing embryos are preseni 
in i brood pouch formed by the expansion of 
the distel end of the oviduct ar the postero- 
dorsal corner of the thorax. The ovary is pre- 
sent about halfway down the posterior abdo- 
men anda single series of pyrifarm testis lobes 
attached to a single duct are present behind the 
ovary. Larvac have the usual three median 
suckers with three ampullae between the 
suckers and many small ampullary vesicles in 
two rows from euch Jateral line as described 
previously for specimens from Western Aats- 
Iralia (see Kott 1963), 
Remarks: The species is closely related to Apl- 
iin phertax (Michaelsen) from New Zeo- 
land, which has a similar number of fine longi- 
tudinal muscle bands, and stomach folds, and 
alsa has. a brood pouch, Consequently, there 
has been some confusion hetween these species. 
Unfortunately, Michaelsen (1924) did not des- 
crthe larvac from his species. Aplidiam plici- 
fern (see Kott 1963) from Western Australia 
has smaller zooids (therax and abdomen to- 
gether about | mm long. posterior abdomen 
2mm) and are densely distributed in the test, 
largely obscuring the systems. In Apliditen 
Pharvay (see Kott 1963) trom eastern Australia 


PATRICIA KOTT 


Figs. 13-17. 


Figs. 18, 19, 


Fig, 20. 
Figs. 21, 22. 


Fig. 23- 


Ritlerella herdmania, (Port Noarlunga). Fig. 13—Young zooid, contracted thorax, Fig. 
14.—Extended thorax of young zooid, Fig. 15.—Zooid with contracted thorax showing 
parastigmatic vessels, Fig. 16.—Thorax of more mature zooid showing parastigmatic 
vessels successively subdividing rows of stigmata. Fig. {7:—Portion of colony. 
Pseudodistoma ceream (Nora Creina). Fig, 18,—Outline of colony. Fig. 19.—Zooid 
with brood pounch, 

Aplidium pliciferum, (Hallett Cove, 8 m). Zooid. 

A plidiumn coleltnlih, (Tapley Shoal, off Trowbridge Light, 17 m). Fig. 21.—Colony. Fig. 
22.—Zooid. 

Synoicium papilliferum. (West 1., sheltered coast, 3 m). Zooid (showing muscles on 
thorax only). 


ASCIDIANS OF SOUTII AWSTRALTA is 


and the Pacific. the larger zooids (thorax and 
abdomen together 3.5 mm long, and posterior 
ahdomen 1.5 mm long) are arranged in cit- 
cular systems. sometumes extending inta more 
elongate and double row systems. radiating 
from the common cloacal openings. In all 
Kott's (1963) specimens the test is gelatinous 
and semi-transparent with red-purple spherical 
piement eclls, and the larvae provide the main 
distinguishing character between the two spe- 
cies. A, phortaxy has larvae with a limited 
number of ampullary vesicles: and a complete 
absence of median ampullae, while the larvae 
of A. pliciferum retain median papillae and 
have many small ampullury vesicles from the 
lateral lines either side of the three median 
suckers. Millar (1966) described specimens 
from Port Phillip Bay as 4. phortax. He points 
out that A, phertax (see in Kott 1963). is not 
apparently the sam species as his colonies 
although he cun only distinguish them by the 
different larval form. He apparently over- 
looked the similarity in the size and form of 
the larvae of bis specimen andl of A. pficiferwmn 
(Redikorvev); Tokioka 1953a: Kott 1963; and 
based his identification ot the ratia of length 
to depth of the hirvae of Michaelsen’s species 
and his own specimens from Port Phillip Bay. 
However, Kott (1963) has already indicated 
thit larvae of A. pkortax (Brewin 19463 trom 
New Zeulund, do have the same rounded form 
as the larvae of specimens of A. phartax (Kott 
1963) from eystern Australia. Tt is apparent. 
therefore, that specimens from Port Phillip Bav 
were erroneously identified by Millar. 


The adul! zooids can definitely be distin- 
gttishe? hy the longer posterior abdomen, the 
smaller size, and the greater crowding of zooids 
of A. plicijerunt 


The specimen from Hallett Cove was taken 
with a specimen of Divtaplia viridis in which 
the zooids wre the same orange colour, The 
specimen from Troubridge Show! was taken 
from 4 spiny crab. 


Aplidium rubricollum Kott, 1963: 103. 
New Record: Upper St. Vincent Gulf. Pre- 
vlous Records: W. Aust. (Rottnest T.).  S, 
Aust; (Reevesby 1}. Vic, {Balnarring 
Beach)—Kott 1963, 


Description: The single colonv is flattened, 
thout OF em thick aiid 3.5 cm in maximum 
diameter, The honiers of the colony are 


rounded. Sand is present basally and some is 
enclosed in the common test but the surface is 
smoath and without sand. The common 
eloucul apertures with Frilled und protuberant 
lips ure present on the surface of the colony 
about 0.3 cm from one another. Spherical pig- 
ment. cells are present in the test and zovids 
show as clear points between the pigmented 
test. In this preserved specimen the pigment 
cells are pale pink. Zooids are small. up to: 
2 mm tong. There are 10 longitudinal 
thoracic muscles. A short pointed atrial lon- 
guet arises from the dorsal surface just anterior 
to the atrial opening which is cenerally on a 
short protuberamt siphon surrounded by u cir 
cular sphincter muscle, There are 1) rows af 
6-8 stigmata, and 4 stomuch folds. 


Remurks: The species is distinguished by the 
form of the atrial aperture and lip. by the nar- 
Tow branchial suc with relatively few stigmata 
in each row and by the body musculature and 
stomach folds, [n the present specimen the 
test is not so thickly invested with sand as pre- 
viously described for this species 


Aplidiam colelluides (Herdman), Millar, 1962; 
125, 


Amarovciam colvlluides Herdoian. 1886: 223, 
New Recenid: OF Troubtidge T. Previeny 
Records: South Afries (Cape af Gond Hopel 
—Herdman 1886; Millur 1942. 


FIGS. 21. 22 


Veseripiion: Rounded velatinous heads on a 
long hard stalk, The head is up to 4 em in 
ength and 2.cm in diameter, The stalk, up to 
20 em in Jength, is hardened by dense sand 
inclusion in the surface test which fades out in 
the test of the head region. ‘lhe stalk i4 
branched basally inta short toot-like processes. 
7ooids are minute, opening around the surface 
of the head. Long thread-like posterior 
abdomina criss-cross in the centre of the head 
and sometimes extend down jnto the stalk. 
Some common clowew! apertures are evident 
around the head and some longitudinal cloacal 
canals were identified, although the form of 
Ihe systems is obscure and difficult ta distin- 
fuish. The thorax and abdomen are of equal 
length and together measure only about 145 
mm, The long, thresd-like post-abdomen is at 
least four times the combined length of the 
thorax and abdomen ‘Where are abour 6 deli- 
cate longitudinal muscles on the thorax. The 


My PATRICIA KOTT 


branvhial lubes are cistinet and rounded. The 
utrial aperture is sometimes produced un a 
fairly long cylindrical siphon but in another 
colony is sessile, the upper border of the atrial 
aperture produced into a pointed languct, There 
are 18 rows of phour 10 short oval stigmuti. 
The oesophagus is long, the stomach is present 
halfway down the abdomen und bas 13 very 
distinct folds. The gonads are not developed 
in these speciniens and it is nut known ta what 
extent they fill the long posterior abdomen in 
muture zooids, 

Remarks: This is the only species of A plidium 
known with a Jong stolk. The size and form of 
the colony, the size-of zootds and their arrange- 
ments. in the present colony are identical with 
the South Africun specimens previously des- 
cribed, The delicate longitudinal thoracic 
muscles and the stomach folds are similar 
The present specimens differ from those des- 
cribed from South Africa only in the larger 
number of rows of stigmata. This does not 
represent a sufficient difference on which to 
establish a new species and in view of the great 
similarity in most characters the specimens 
probably represent one species with a wide cir- 
cumpolar distribution in the southern colu- 
temperate region. 


Synviciuin = papilliferum = (Michaelsen), 
1963; 87, Millar, 19A6: 360. 
Macrollinumn pepilliferion Michuelsen, 1930: 
520, 


Katt, 


New Records: Port Noarlunga reel, West 1. 

(neur Penguin Rock), Previony Records: 

W, Aust. (Bunbury to Nornalup)—Michael- 

sen 1930; Kotr 1963. Vie. (Nepean Penin- 

sula)—Millar 1966, The species is known 

intertidally and to LS m, 

FIG, 23 

Deyeription: Un life the colony is dark red or 
bright brick red. Flat-topped to rounded colo- 
nies, narrowing basally Wa common stalk or 
encrusting. Zooids Jie parallel i the test and 
open on the upper surfiuce. ‘Vhe colony is firm, 
gelatinous. There are circular systems arcune 
protuberant common cloacal apertures. The 
branchial aperture has 6 small pointed lobes 
and there is a small circular sphincter muscle 
at the base of the branchial siphon. The atrial 
aperture ix opposite the first to second row of 
stigmata. It is surrounded by a well developed 
circular sphincter muscle. and is extended into 
w short cylindrical siphon. The anterior boner 
ol the atrial aperture is produced into a long 
mnscular lip, broken into 3-4 minute pointed 


lobes terminally, There ure !0 very fine longi- 
tudinal muscle bands on the thorax which is 
very delicate and transparent. There are 
{0-12 rows of about 10) stigmata in each row, 
Che hody wall below the atrial aperture is pro- 


duced into the small rounded pupillae charac- 


teristic of Synoiciun spp. The wall of the 
slomach is riised into faint mulberry-like swel- 
lings. The posterior abdomen is shart and 
there is no constriction between it and the 
abdomen, 

Rentarks: Both colony and zoaids conform 
with previous descriptions in all characters 
except the reduced mumber of rows of stig- 
mata. The species has been recorded fram 
south-western Australis along the south coast 
of Australta ta the Nepean Peninsula in Vice 
toria (Millar 1966). 


Tamily DIDEMNIDAE 
? ‘Trididemunm spiculatums Kon, 1962: 281. 


New Record: West 1. (near Penguin Rock), 
Previous Records: W. Aust. (Rottnest 1., 
Point Peron), S. Aust. (Quter Harbour) 
Tas. (Wreck Bay). Qld. (Heron b.I— Kare 
1962, 
Description; Living colonies pile pink, enctitst- 
ing. Small, almost spherical spicules with up 
ty 12 points in optical transverse section, 
evenly distributed throughout the test, and 
oeeasonally large spicules with fewer rays. 
There are small thoracic common cloacal cavi- 
Ges. Zooids are smull with three rows of stig- 
mata, The atrial aperture is wide. cxposing a 
lurge part of the branchial suc. Gonads are 
nut mature in the present specimens, 
Remarks: Colonies generally conform with 
specimens previously assigned to this specics, 
ulthough the proportion of smaller isner-lke 
spicules lo larger stellate spicules with about & 
rays in optical section. is greater in the present 
specimen, Colonies with mature zdutds are 
desirable for positive identification. 


Leptoclinides rufus (Sluiter). Tokioka, 1952; 
92. Kott. 1962: 286 and synonymy. 
Eldredge, 1967: 221. 

Rossenpranate refus Sluiter, $9092 72; 1913: 
New Records: Off Pore Gawier, Hallett 
Cove, Port Noarlunga, Rapid Head, West T., 
Wright F, Previous Records; §, Aust. (Port 
Noarlunga). View (Shoreham). Tas, (Maurin 
1}. NSW, (Port Jackson)—Kott 1962. 
Old. (Heron T.)—Hustings 1931. New Zea- 
land {Great Barrier t.. L. ¢luiteri}—-Brewin 
1950b; (?Stewart I, L. nNovdezelatidiae}— 


ASCIDIANS OF SOLTH ALSTRALIA 7 


Brewin 1958a, (?Chutham Rise, Lauran 
ticis)—Brewin 1956: (North t.)—Michgel- 
sen 1924) Brewin 1958b; Millar 1960, Indo- 
Pacific fArafura Sea, Indonesia, Hawati}— 
Tokioku 1952; Stuiter 1909; Eldredge 1947, 
‘The spectes is Known intertidally and to 36 
m (Sluiter 1909). 


Deseripiion; Encrusting colonies, Living 
Specimens: while matrit with grey oar dark 
animals, or orange ta light fawn (Port Noar- 
lunga); or dark reddish brown folf Hallett 
Cove), mottled white to uniform light grey 
colour (Wright 1.).. In preservative all calonies 
ire While to orange-white or streaked and 
blotched with grey. ‘Vhe colonies are investing, 
sometimes extensive. Cloacal cavities radiate 
from randomly distributed apertures, Zoids 
ire sametimes present in the roof of the com- 
mon cloacal cavity. Spicules ure present in the 
surface tese but basally the testis jelly-like and 
transparent. There are 9 tongituclinal muscles 
ev the thorax. The posteriorly directed atrial 
siphon has a wide circular sphincter muscle. 
There ure 4 rows of 10 to 12 stigmata, There 
is a superficial layer of bladder cells and small 
oval to spherical pigment cells are present 
amongst the surface Jayer of spicules. A literal 
organ is present opposite the middle of the 
fourth row of stigmata, Cloncal apertures are 
present, espectully uround the burders of the 
calony. Canals. at thoracic level radiate from 
the cloacal apertures between clumps of zooids 
although sometimes they extend deeper to 
abdominal level. The cloucal canals around 
the border of the colony ire often completely 
sub-abdominal. The ypicules are of the usual 
stellate form, 0,0/-0.04 mm in diameter. 
Larvae are present in some colonies from 
Hullett Cove. They are of usual form, fairly 
deep with 4 paired ampullue. In one colony 
from Hallett Cove (dark reddish brown in life) 
no common cloacal cavities were present and 
zooids Were not mature, nor were zooid open- 
ings to the exterior detected, The arrangement 
of spicules is characteristic of this species and 
jl a8 probable that the colony is one in which 
sexual reproduction ts completed and new vege- 
tative buils are developing. 


Remarks: The species ix distinguished by the 
complete absence of spicules from the basal 
layer of the test, sometimes giving the colony 
a very fleshy appearance. The characteristic 
common cloacal system and the distinct mus- 
culature on the thorax, together with the pos- 
tetiorly directed atrial siphon and the spherical 
to oval pigment cells are distinctive, 


Leptoclinides kingi Michwelscn. 
Palveyneraron aibiint. Van Nate, (STB: b55, 
Hartmeyer, 1819: 136, 
Leplectinidis dublax f. kinet 
(930: 507, Kou. 1942) 2489 
New Record; Upper St. Vincent Gulf. Pre- 
vious Records: W. Aust. (Fremantle, Albany) 
—Michaelsen 1930. Old. (Sarlna)~ -Kott 


Michaelser, 


1962, Philippines (dolu Light} —Van Name 
1918. The species is known intertidully anu 
to 18 m. 


FIGS, 24, 25 


Deseripnion: Vhe colony is massive with the 
surface tuised into mounds und single cloacal 
apertures ut the apex of each mound. Each 
mound ts lormed by thickened basal test often 
with embedded pirasites, Zooids are present 
in the surface test above the very »xtensive 
posterior abdominu!l spaces round the centre 
of each lobe or mound, The zooids ure large 
with 4 rows of about 12 stigmata, There are $ 
very fine longitudinal muscles on the thuran, 
The ‘spicules are very small, (1.01 to 0,02, mm. 
and are ranged in a shallow laver at the level 
of the branchial siphons. They are only very 
sparse elsewherd in the test. There is 4 surface 
layer of bladder cells. 


Remarky: The elevation of the surface of this 
coluny into mounds or lobes with terminal 
common cloacal apertures characterises this 
species, which was previously regarded as a 
form of Leptoclinides dubius (Sluiter). Lepte- 
clinédes dubius is distinguished from ihe present 
specics hy its larger spicules and by the 
utrangement of common cloacal system with 
openings around the margins of each opley, 
us in B, rufus. In £. kingi large cloacal sys- 
tems with terminal openings develop fram the 
centre of the colany, As both forms have been 
recorded more of less ever the same geographic 
unge it as unlikely that they represent geo- 
graphic subspecies of the ons species, and in 
view of the different development of the com- 
mon coacal systems it is probable that they rep- 
resent different species. The ony gut loop 
which is bent anteriorly to Form a double loop 
is a character shared with Leproclinides didits, 
Posteriorly directed atrial siphons of the zooids 
open inte the common cloacal cavities and 
canals, The openings sometimes appear 5 
lobed due to the arrangement of spicules 
around the aperture, The genus Askonides 
Kott, 1962, therefore cannot be distinguished 
from Leptociinides and A. imperfectur and A, 
ceelenterwius are distinguished from other spe- 
cies of Leptoclinides only by the extent to 


1k PATRICIA KOTT 


which zooids open directly into Ihe common 
cloacal chamber rather than into cloacal canals. 
Their relations are set out in the following key: 


|, Single systems develop around central com- 
mon cloacal cavities with terminal epen- 
ings Passeeccietge ie Sade 

1, Numerous systems develop around peri- 
phery of colony 3 

2 Spicules accumulated in surface layer o 
test; spicules 0.0)-0.02: larvae with 4 
paired ampullae; most zoaids open into 
cloacal carvals L. kingi 
2. Spicules throughout; spicules 0.04-0.08, 
lurve with reduced ampullae; most 
zooids open direct into common cloacal 
cavily L, covlenreranus 
and L. imperfectius 

3, Spicitles 0.01-0.02; double gut loop... 
L. dubius 
3. Spicules 0,02-0.04; simple gut loop 

L. rufus 


Leptoclinides reticulatus (Siuiter}. Kott, 1962: 

285 and synonymy. 

Didemniun refientaive Siuiter, 09> ha. 
New Record: Tipata Reel. Previous 
Reiords; Qld. (Noosa to Mackay, Heron 1, 
Low Is.)—Hastings 1931; Kolt 1962, New 
Zealand (North 1.)—Michaelsen 1924, 
Japan—Oka 1927; Tokiokn 1953a, 1953b. 
Indonesit —Stuiler 1909, 7Philippines-Van 
Name 1918. Indian Ocean (Ceylon) —Herd- 
man 1906. 


FIG. 26 


Deseriplion; Young colonies were taken invest- 
ing Micrecosmus squarmiger and Pytire irregu- 
lariy. Frequent common cloacal openings are 
scattered over Vhe surface. There is super- 
ficial layer of bladder cells with orange and 
black pigment in stellate cells forming streaks 
on the surface. Spicules are present benenth 
this superficial layer and are reduced in den- 
sity toward the base af he colony. The spi- 
pules ire sicilate with about 7 conical rays in 
optical transverse section and from 0,03 to 
O05 mm ty optical section, 

The vrimary clovcal canals are deep, but in 

these specimens. do not extend posterior to the 
zooids. The znoids are small with the usuul 
4 rows of stigmata and a large posteriorly 
divected atrial siphon. There are 4 testis lobes 
and 44 coils of the vas deferens. 
Remarks! This 1s the most southerly record for 
this: conspicuous. and widespread species, dis- 
tinguished by its unique stellate pigment cells 
whicl) form the charucteristic “ogertike” mark- 
ings on the surface, 


Didemnaum tambitum (Stuiter), Kou, 1962: 317 

and synonymy, 1971: 19%. 

Didemnoides lanbitem Sluiter, 1900, 18, 
New Record: Aldinga “drop off. Previous 
Rerords: N.S.W.---Kait 1954, 1962, New 
Zealand (Chatham I.. North L.. South [,)— 
Sluiter (1900; Michaelsen 1924; Kott 1971: 
and unpublished records from Otage (call 
R. Ceump) and Stewart I, (coll. . Barham). 


Description: Two clavate lobes arise from a 
common base, Maximum diameter 1.5 em 
and maximum height 3.0 em. There are traces 
of orange pigmen{ in the surface Lest, but no 
superficial layer of bladder eclls, There is a 
layer of spicules in the surfuece lest which 
ceases abruptly at ocsophugeal level. Thin 
Jayers of spicules line the common eloacal 
canal. Spicules. are absent at the abdominal 
level of the zooids, and in dhe centtul test core. 
They are 0.01 to 0.05 mm and stellate, Ter- 
minal cloycal aperture opens into the charac- 
(éristic common cloacal cavity surrounding the 
central core of test. Zooids are small and 
crowded in the surface layer of test, “The atrial 
aperture is wide and open. There are 8! coils 
of the vas deferens around a single testis lobe. 


Didemnum patulum (Herdman), 

Leptaclinym petuiim Herdman, 1499: 92. 
New Record: Aldinga, Previous Recerels: 
Vie, (Port Phillip Bay)}—unpublished regord 
N.S.W. (Port Jackson) Herdman 1894. 

FIG. 27 
Deveriplions Tough, investing colonies. In 
preservative the specimens ure white with grey 
streaks and blotches formed by palehes of stel- 
late pigment cells in the surface test, especially 
in the région of the commun cloacal canals 
The surface of the colony is marked off into 
alightly raised rounded arews where solid pil- 
lars of test traverse the coramuan cloacal cavity, 
Zooids are embedded! in the periphery of these 
pillars of test and open to the surface around 
the raised ares, “Whe cloacal cavity is thoracic. 
The surface layer of test is especially thick and 
the zooids huve especially long and musculus 
branchial siphons which estend through this 
surface layer of test. Spicules often form a 
plug inside the branchial siphon—possibly 
caused when the superficial layer of test is 
pulled down into the aperture as it is Teiracted 
into the surface of the test. The branchial 
siphon is almost the same length as the rest of 
the thorax. The atrial opening is wide, expos- 
ing a ptt of the dorsal surface of the branchial 
sue, The anterior border of the atrial opening 


ASCIDIANS OF SOUTH AUSTRALIA iW 


\§ produced into a natraw pointed languer, 
sometinves bidentate at the tip. There ane con- 
spicuous circular muscles in the branchial 
siphon, in addition to the usual longitudinal 
muscles that extend down the length of the 
thorax and into the test to form a short retrace 
tor muscle. The abdomen, of the usual form 
lor this genus, is especially small. Oesophageal 
buds are present but the gonads are nat mature, 


Remarks: The grey veins in the surface identify 
This specimen with Herdmun’s species, The 
long branchial siphon and atrial lip wre also 
distinctive. The specres is especially common 
in Port PRillip Bay, but is not common in St. 
Vincent Gulf The species also strongly 
resembles 2, tabulate Sluiter from the East 
Indies and Aru 1. (sec Sluiter 1913; Kott 1962). 


Didemoum moseleyi (Herdman). Vat Name, 
1918: ISL. Tokioka, 195Sa: 212: }955h; 
44; 1959: 226; 1961: 106. Katt, 1957b; 
136: 1962: 328 and synenymy, Eldredge. 
1967; 212, 

Leproctinuen moseleve Merdman, 1&6: 272. 

fupreclinion incanim Hendman, (898° 90, 

Herdman & Riddell, (813; 888. 
New Records: Cioose |, Carickulinga Head. 
West 1. Previous Records: W, Aust, (Rote 
nest L., Point Peron, Trigg 1). S. Aust. 
(Reevesby LL). Vic. (Balnarring Beach )— 
Katt 1962. Tas. (Spring Bay, Maria I.}. 
N.S,W, (Port Jackson, Port Stephens, Coffs 
Hatbour)—Herdman 1899: Kott 962. 
Indiun Ocean {Southern Arabin}—Kott 
19S7h. Indonesia (Arafura Sea)—Sluiter 
1914, 1913; Tokioka 19553. Pacific Ocean 
(Patau is. New Caledonia, Philippines. 
Hawallan ts, Marshall Is.) —Herdman 
1886: Van Name 1918; ‘Tokioka (9556, 
1961; Eldredye 1967. 


FIG, 28 


Description: Investing sheets, There is a very 
thin Jayee of surface test which is often tatged 
imo spicule-filled conical papillac between the 
branchial apertures, The cloacal cavity is 
thoracic and the thoraces of szooids are 
enclosed in an independant test sheath, The 
atrial opening is wide, in all cases exposing the 
branchial sac tothe cloacal canal, Spicules are 
0,02 to 0.04 mm in diameter with no more than 
10 pointed rays in optical transverse section 
and are densely distributed throughout. Zooids 
are colourless. They are minute. the branchial 
sac espycially small with four rows of only & 
stigmata, The yas deferens coils 64 times 
around a single undivided testis follicle, In the 


specimens from West 1. and Carickalingy Head 
there is a smull laters] organ opposite the last 
two TOoWs of stigmata. 


Remurks: Eldredge (1967), discussing the 
difficulties in distinguishing between the present 
species and D. candidun, hus suggested that in 
2, candidum the surface test is always smooth. 
the atrial aperture is w small slit and Jateral 
organs. are always absent. He has not been 
able to confirm the presence of larger numbers 
of yas deferens coils for D. cenedidum (Kott 
1962) nor is the condition of any of these 
characters constant in specimens previously 
ascribed to the species, Only the regularly 
stellate spicules and dark pigmented zooids of 
the present specimens uppear to distinguish 
them from D. movelevi which has a variety of 
diferent types of spicules. 


Didemnum candidum Savigny, 18i6; |94, 
Michaelsen, 1924; 358 und synonymy. 
Van Name, 1945; 83, Hastings, 1931: 94, 
Brewin, 19467 98; 1950u: 55: 1950b; 
345; 1951: 104: 1952b: 188; 1956: | 22; 
IYST= S77, 196 14. Tokioka, 1954y; 
246; 195Sa: 45. Koti, 1954: 162; 1962: 
327. Eldredge, 1967: 23. 

The above synonymy refers only to Indo- 
Pacific records. For full jist of synonyms 
see Fldredge 1967; 213. 


New Records; West T., Wright lL, Previnviy 
Records: South-western Australia, Tasmania, 
north-eastern Australia. the Envlish Channel, 
Trish Sea, West Africa, South Africa and 
East Africa, Red Sea, Mediterranean Sen, 
New Zealand. west and mid-Pucific Ovenn 
(Marshall is. and Hawaiian ts ), the Carib- 
bean and West Indics and the est coast of 
the USA. Records are lacking from the 
north Pacific and west coast of the Amenecan 
continent, but elsewhere the species pevurs 
widely in temperate and tropical regions. 


FIGS, 29, 40 


Deseription: Colonies arc Hat and investing, 
small and rounded or more extensive sheets. 
The test has dense spicules throughaut, In 
preservative the zovids are brown and show 
through the white spicules. The common 
cloacal cavity is thoracic but extensive and 
limited only. by thin layers of surface and 
slightly thicker basal test in which the abdo- 
mina of the zooids are embedded, Thoraces 
cross the common cloacal cavity in an indepen- 
dent sheath of test, Spicules are dense through- 
out. They are 0.02 to 0.03 mm in diameter 


20 PATRICIA KOTT 


and demonstrate the same range in form pre-almost cylindrical marginal rims stiffened by 
viously described for this species with up to 1Sthe dense spicules enclosed in the test. Zooids 
or more rays in optical transverse section. Con-are very small. There are 4 rows of about 8 
spicuous common cloacal apertures present onstigmata. No gonads were distinguished in the 
the surface are surrounded by protuberant,present colonies. 


ASCIDIANS OF SOUTH AUSTRALIA 2] 


Remarks; The present colonial systems are 
typical of the species although no gonads 
Appeared to be mature, It was not possible to 
confirm Eldredge's observations concerning the 
slitlike atrial opening as in the extended 
zouus of the present colonies, these were wide 
open, exposing a great part of the dorsal aspect 
of the branchial sac. The vartety of spicules. 
therefore, rémain the principal distinguishing 
character for this species. Curliste (1954) has 
characterised specimens of 2D. candida 
Savignay Fram the North Sea, the English Chan- 
nel, north-west Afvicu, the Medilerrancan and 
the Red Sea (type bocality) by the absence of 
the third adhesive pupilla in the lurvac, and 
Lafargue (1968) confirms the condition for 
specimens trom. the French coast. The speci- 
mens agree in all other respects with those des- 
cribed from New Zealand, Australia, Malaysia, 
Japan and the Atlantic coast of America. 
Carlisle concludes, therefore, that: “D. cendj- 
dum is a tropical and lemperate species extend- 
ing from the West Indies to the East Indies, 
New Zealand and Japan”. 

However, later workers have not observed 
the universal absence of a third adhesive papilla 
in luryae from these localities, while there are 
the usual three larval papillae in Australian, 
New Zealand and Japanese specimens. It is 
possible, therefore. that two separate specics 
are Involved. 


Polvsyncraton orbiculum Kott, 1962; 300. 
New Record; Rapid Head. Previous 
Recordy: W, Aust, (Rottnest 1). 8. Aust. 
(Por Noarlungy)—Kott 1962. 

Peseripion: The preserved colony is light 

pinkish brown, owing to the darkly pigrnented 

vooids seen through the single layer of spicules 
presemt in the thin. surface test. The dark 
coloured zooids are ulyo seen through the 
branchial Openings clearly marked on the sur- 
face test. There aro the usual vesicular cells 
arranged in « complete circle around the 
branchial openings. and interrupting the ather- 
wise even disuibution of the spicules in the 
surface test. There is an extensive thoracic 


cloacal cavity, crossed hy the thoraces of the 
zoulds, exch with a discrete ventral sheath of 
test. There is a iitteral organ about halfway 
down the thoracic test sheath, The vooids are 
small, with 4 rows of stigmata, There is a long 
retractor muscle, These specimens conform 
with those described previously (Kott 1962} 
in all respects; however, the gonuls are not 
muture mn the present zovids. 

Remarks, The condition of the cloucal cavity, 
the dark pigmented zooids, the rather Jarge stel- 
late spicules and the unique, large transparent 
vesicles in regular circles in the surface, 
together, charucterise the species. 


Echinoclimom $yerrilli Van Name, 1902: 372, 
Kot. 1962: 312 and synonyms, 
Diplosoma VLissoclinum) verrillt, Eldredge. 
1967: 242, 

New Recards: Hallett Cove. The species 
has been observed investing the underside of 
rocks aj 4 depth of S20 im at many Joca- 
trons in St. Vineent Gulf where conditions 
are quiet. The colonics are so fragile, how- 
ever, that they usually break up when 
removed (S. Shepherd, pers. comm.) _Pre- 
vious Recares: Vas, (West Coast)—Kot 
1954. America (West Indies, Florida)— 
Van Name 1902, 1945; Hartmeyer, 19f9— 
11: Plough & Jones 1937. Africa (Accra) 
—Millur 1953. Japan {Sagami Bay)—- 
Tokioka 1958. 
FIGS. 31-35 

Deveription: Living colony soft. white, jelly- 

like. In preservative the present colony is deli- 

cate and soft, It appears to be investing but is, 
unfortunately, damaged and its exact form 
could not he determined. Spicules are mostly 

G-rayed, but there are alse spicules with 4 anu 

with 3 rays. They form a dense spiny, tough 

capsule around the abdomina of the zaojds but 
are sparse in the remainder of the cobony- 

Zooids ate arranged more or less in the dauble 

rows previously described (Van Name 1945) 

although common cloacal openings were not 

deiected. The cloacal canals spread usit 
beneath the zonids whith are retained m the 


Figs. 24, 24, beptoctinides kingi, (Upper St. Vincent Gulf, 10-12 m), Fig. 214.—Spiciles. Fig. 25. 


Thorax, diagrammatic, showing mus 


Gut loap, 
Fig. 26. Leptoclinides: reticulatus, (West 1,, under boulder), Snicules, 
Fig. 27, Didemnum parnlam, (Aldinga “drop-off, 3-8 m). 

cutature. 
Fig, 28 Didermnum moseleyi, (Cavickalings Head, 5-6 m), Spicules. 


Figs, 29, 30, Bidemunm vandidum, (Wright T. 
Fig 30—Spicules, 
Figs, 31-35. 


rough coast, 10 m), Fig, 28—Diayram of colony 


Echinoellnum verilli. (Hallet Cove, 8 m), Fig. 31. Spicules. Figs. 32, 33, 34.—Larvae 


of incyeagiog maturily, Fig. 35,—Matare anterior ampullae of larvac. 


ps PATRICIA KOTT 


surface test, Zooids ure small with large 
lateral organs on each side of the thoras. 

Larvae fre large with a short tail which. 
when extended, is only hall the total length of 
the lurva. There is a large ocellus and an -ote- 
fith, At jeast one precocious bud is present 
although the exact number is obscured by the 
laver of spherical to oval granulate bodies that 
extend around the posterior half of the body of 
the lurva. 


Antenorly there ure the usual three adhesive 
papillae in the median tine and [4 ampullae 
from the Imeral lines on either side of the 
suckers. Ipitially these lateral ampullae are 
very small and sessile. Subsequently they 
jncveuse in size und become “tear-drop” in 
shape supported by very narrow stalks from 
we lateral line. 


Remarks Wis unfortunate that the present 
eolony is so damaged thal its shape cannot be 
discerned. Although previously deseribed 
specimens have been clavate (Kott 1954) Van 
Name [945] the present damaged colony ts 
investing and living colanjes have been 
observed investing the under-surface of rocks, 
It is possible therefore that two distinet species 
may be involved, characterised by a difference 
uy the consistency of the text and in the shape 
of the colony. 


‘The soft nature of the colony and its ten- 
dency to break up hus probably been the cause 
of the lack of records of this form, which is 
reported as common in St. Vincent Gulf- 


Eldredge (1967) has suggested that the 
genus. is synonymous with Diplexoma (Lisin- 
clini), due to the similarity of the cloucal 
systems avid the fact that tetrahedral spicules 
ure not unique in the family Didemnidac. 
Eldredve's contention cannet he maintained, 
Vhe common cloucal cavity in the two genera 
is extensive and extends posterior to the zooitts 
which remain connected to the hasul test by 
strinds of test. However, the cloacal syster 
in Echinocliniwn differs from that in Diployoma 
(Livsoelinimy in the absence ol the secondary 
cloacal spaces around the thoraces of the zaoids 
which remain connected to and in the surface 
test in continuous rows. In Diplosema” { Lice. 
clinuar) the secondary cloacal spaces sepacate 
cither the thoraces, or the whole zooids, from 
one another. Further, the spicules in Eehine- 
elinum are very much larger (0,05—0.1 mm) 
than those generally found in other genera of 
the famlly and, in addition co their unusuul 
form and size, their distribution in the coluny 


differs. entirely from other venera of the 
Didemnidac. The capsules formed around the 
zooids by ihe spicules are reminiscent of the 
eapsules formed in Cystodyres spp. and in no 
other venus of the Didemnidue do the spicules 
remain in such an intimate relationship with 
the zon. 

The genus is further distinguished by a 
Unigue larval form with wv multipleity of 
narrow-slulked epidermal ampullae and prece- 
cious. buds. The larvae of P, aspiculatum and 
D, (Lissoclinnin) spp. show a similiar oirkedt 
increase in the number of lateral ampulbic, The 
ampullue in Eehinoctnum wre unique, how- 
ever, in their distinet “tear-drop” shape, their 
natrow stalks and their diserete origin from the 
lateral jine without subsequent stibdivision. 
Precocious budding generally occurs in the 
farvae of Diplosema spp. and in D. Lésxa- 
clinum) spp. However. it alsa occurs in 
Didemaunt (B, psendodiplasamu— Rott 1962, 
and D. ternarenuni—Koie L866) and in Paly- 
syecraton (P. aspiculatim—Koir 1962) so can- 
not be considered characteristic of any single 
genus. 

The zeunular bodies present in the larval test 
are indeed similar to those found in O. 1 Livsa- 
clinum | fragile—Eldredge 1967 and D. | Lisso- 
clingeny) dstrearinmn—-Kort 1962, They do not 
like up huematoxylin stains (Eldredge 19671 
and thus de not appear to be calcareous spi- 
cules nor their precursors. as Kott (1962) had 
sutycsicd. However, despite che relationship 
with D. ( Livsoclinunty medicated by these 
enclosed granules. the genus is distinct from 
ether genera in the Didemnidae und entirely 
justifies. iis Laxonomic position as a monotypic 
genus in that family- 


Didemnunt sp. 

Renard: West I, (near Penguin Rock) 
Deseription: Living colony “yellow, crusuase". 
In preservative the investing colony is a light 
fawn colour. There are common cloacil aper- 
tures with large spicule-filled lips scattered over 
the surfuce of the enlony. Zooids are suspended 
between the basal and surface layers of tesr 
by connecting columns of test in which the 
abdomina are embedded in clumps, although 
the vooids are separated from one another in 
their own discrete sheath of Lest, open to the 
vommon cloacal cavity on the dorsum. Stellate 
spicules are thick throughout the test. The 
branchial siphons are Carly tong with distinct 
circular muscles. There are Jatge oval hitecul 
organs on either side of the thorax. There are 
four rows of stigmata. 


ASCIDIANS OF SOUTH AUSTRALIA 


Reniarky! The gonads are not developed and 
a definitive identification of the genus is there- 
fore not possible. The cemditiun of the colony 
with a well developed posterior abdominal 
cloacal canal is reminiscent of cerluin species 
Of Didemnrium, 


Suborder PHLEBOBRANCHIA 
Family CORELLIDAE 
Subfamily rilencsomaTiNar 


Khadosoma tireicuns (Savigny). Kotr, 1952: 
417 and synonyity.. Tokioka, 1952: 111; 
W953q: 230, 

Phallisia vcica Saigny, 1B162 102, 

Rhadoxoma papillosum. Van Name, 1918: 

113 and synonymy. Hartneyer. 1919: 99, 
New Record: Hallett Cove, — Previons 
Reeords: NW. Aust. (Cape Jaubert) —Harst- 
meyer 1919. S$. Aust. (Port Nourluinga)- 
Old.—Kon 1952. Indonesia—Sluiter 1904; 
(Aralura Sea)—Tokioka 1952, Indian OQcean 
(CeylonJ—Herdman 1906, Pacific Ocean 
(Philippines. Califarnia)—Wan Name 1918, 


1945; (Chile}—Traustedt 882, A835: 
(China)—Stimpson 1855;  {Japan)—Oka 
1927: Hartmeyer 1906; ‘Tokioka 1953a. 


Red Sea—Ehrenberg 1828, Mediterranean 
—tLacaze-Duihiers 1865. The species is also 
recorded From the Caribbean region (Van 
Name 1945) 


Remarks; Nothing further cun be added to the 
deseriptian of this cosmopolitan but rare 
species. His never taken in large numbers, 
nor is it taken very often, The species is, 
however, not inconspicuous, It is probable 
that, with its highly developed closing mecha- 
nism, it may exhibit a high degree of vivipary. 
In which case it is probable that rehiytively few 
larvae are incubated, and that the free-swim- 
ming time of larvae is shart. “he dispersal of 
lurvae could be. therefore, limited, and the 
survival of the apparently small populations of 
the species enhanced by larval settlement close 
to the parent zooids, The specics has heen 
taken from a wide variety of depths. Lnfor- 
tunately, little is known of the current condi- 
tions at locations from which the species has 
been taken, but it is possible that it favours 
less turbulent conditions where there is misimal 
current flow so that the larvae would be even 
lest exposed to dispersal, 


Only a single specimen ts present in this col- 
lection, 


iz 
“a 


Subfamily COREBLINAE 
Corella cumyota Traustedt, (882; 271, Kot, 
1969; 84 und synonymy; 1971 > 20 
New Records; Hallett Cove, King Beach. 
Previous Records! W. Aust (Trigg 1.j— 
Kott 1952. Vie, (Balnarring Beach, Franks- 


ton) —Kott 1952, Millur 1966, Tas, 
{D'Entrecasteaux Channel), New Zealand 
(North and South 1s.)—Sluiter 1898: 


Michielsen 1922; Brewin 1946, 1948, 1950a. 
1957. 19640. South Africu—-Stuiter 1898; 
Michuelsen 1915; Millar 1955, 1962. ‘The 
species also has i wide circumpolar distribu- 
tion in the Antarctic (Kott 1969). 


FIG, 36 


Deseription; The living $pecinens were noted 
transparent anc’ no colour was recorded. 
There are both separate individuals and indi- 
viduals ageregated together more or less in a 
line. Zooids are generally fixed to one another 
er to the substrate by almost the whole of the 
fight side, The atest is thick, gelatinous and 
semi-transparent. On the right side of the 
body where it is fixed to the substrate the 
boily wall is especially’ thin and there are no 
nuscles except those which radiate a shoarr dis- 
tance from the branchial siphon. On the upper 
or left side of the body there are mostly trans- 
verse muscles branching and ramilying and 
some short and more regular transverse muscles 
in a single row extending, sround the ventral 
border. The branchial siphon is terminal and 
on a short cylindrical siphon. The atrial aper- 
ture is sexsile and from the posterior third ef 
the dorsal border, ‘The branchial sac, gut and 
gonads are of the usual form characteristic of 
the genus. 

Remarks: These specimens do not differ in uny 
Way from other specimens of this ubiquitous 
species which has been recorded in very large 
numbers from open sea locations in ecireum- 
polar waters of the Antarctle and the sub- 
Anturetic (Kot 1969, 1971). The vorthern 
extent of the recorded range is at Trigg I, 
{Kott 1962) on the western coast of Australia, 
hut the species has not been taken on the 
eastern count of the Australian mainland: the 
most easterly record on the Australian cuast is 
at Frankston in Victoria (Millar 1966). 


Family ASCIDIIDAE 


Phallusia depressiuscula (Heller), Kutt, 1472: 
8 and synonymy. 
Ascidia depressiustula Heller, 1878: = Herd- 
man, 1906; 305. 
Ascidia julinéa, Vasseur, 1967; 129. 


am PATRICIA KOLT 


New Records: Tapley Shoal. off Part 
Gawler, off Grange, off West Beach, Hallett 
Cove, off Port Stanvac, Wright I. Previous 
Records: W, Aust. (N.W. Aust. Shark Bay, 
Fremantle)—Hartmever 1919; Michjelsen & 
Hartmeyer 1928; Millar 1963. N.S.W- 
(Port Jacksonj—Herdman 1899. Qld. 
(Great Barrier Reef)—Hastings 1931, Kott 
1952, 1946. Bass Strait (Enst Moncgeur 1.) 
—Herdman 882. Pacific Philippines, 
Palao Is. New Caledonig)—Vun Name 
L913; Tokioka 7250; Vasseur 1967, Indo- 
Malaya (Ceylon, Indonesia, Arafura Sea)— 
Heller 1878: Hetdman 1906; Stuiter 1919: 
Tokioka 1952, The species. is krown inter- 
tidally and to 52 m, 


Description: Living specimens from off Hallete 
Cove are noted as large, white or transparent, 
and cammon on sandy bottom. Many living 
specimens. however. ore bluixh, with black and 
yellaw markings. The preserved specimens 
may be whitish, or blackish grey und may have 
black spots in the surface test, The test is 
thick and firm, smooth on the surface with 
rounded ridges and swellings, The individuals 
reach a large size. ‘The present specimens 
exhibit the range of variation described by Kott 
(1966) for the species, 


Remurks: The relationship of Phallusia julined 
Sluiter lo the present species remains in doubt. 
The specimens in the present collection have 
the atrial aperture from the anterior third of 
the body while specimens of P julines have 
been distinguished by the position of the atrial 
aperture From the posterior third of the body. 


Ascidiq sydneyensis Stimpson (?part), 1885: 
387, Kott, 1972 and synonymy- 


New Records; Taplcey Shoal, Hallett Cave, 
Port Noarlunga, Wright 1, Previous Records: 
W, Aust (Cape Jaubert to: Albany )—Hart 
mever 1919; Michaelsen & Hartmeyer 1928; 
Millar 1963, S, Aust. (Victor Harbor, Port 
Noarlunga). Vic. (Ralnarring Beach, Point 
Leo, Port Phillip Bay}—Kott 1952; Millar’ 
1960; 1963; 1966, Tas. (Spring Bay). 
N.S.\W, (Port Jackson)—Stimpson 1855; 
Herdman 1882, 1899, Qld. (Caloundra to 
Townsville)—Sehmeliz 1879; Koil 1962, 
1966. Indonesia (Arafura. Sca)—Sluiter 
1886, 1904; Tokioka 1952, Pacific Qcean— 
Traustedt 1885; (Palao y., New Caledonia) 
—Tokioky 1950; Vasseur 1967, Japan— 


Harimeyer 1906, Tokioka I9S3a, 1954b. 
Indian Ocean (Seychelles) — Michaelsen 
1918; (Zanzibari—Traustedt & Weltner 
1R94; (East Africu}—Millar 1956. South 
Africa—Heller 1878; WHurtmeyer 19t1, 
19) 3; Shutter 1898: Millar 1955, 1962. The 
species is also recomed from. the Caribbean 
region (Van Nume i435). [tis taken inter- 
tidally and to 31h m, 


FIGS, 37, 38 


Descriprion; Vhe living specimens are. trans- 
parent and fleshy. The largest specimens in 
the present collection are 20 cm long und 12 
em wide. The test is thin, but firm and tough, 
and in larger Specimens slightly leathery. 
There is sometimes. especially on the larger 
specimens, a very sparse encrustation of weed 
and worm tubes. Both the branchial and atrial 
ipertures. ore on short cylindrical siphons and 
ure usually about half the body length distant 
fram one another. Specimens may be fixed ta 
the substrate by the posterior, ventral, or left 
side of the body. The branchial siphon is 
turned away Front the atrial siphon lo varying 
extents. here is a row of short transverse 
muscle bands around the dorsal and ventral 
borders of the right side of the body, The gut 
is always filled with mud, which appears to 
accumulate during the life of the individual 
until in lirger specimens the gut is so swollen 
with mud that the branchial sac is occluded 
und confined ta a small area to the right and 
dorsal to the mud-filled gut, This mud hegins 
to collect, in smaller specimens, in the descend- 
ing limb of the primary gut loop, beyond the 
stomach, and it extends from there into the 
rectum and continues to accumulate in these 
sections Of the intestine. 


Remarks: The physiological significance of the 
mud-filled gut which appears. to be characteris- 
tic of this species is nol known. It has been 
noted in specimens from all parts of the Puci- 
fic. Abbott (pers, comm, 1955) noted that it 
appears to be associated with the termination 
of the typholosole at the top.of the gut swelling 
instead of extending further down the intestine. 
The stomach appears to be free from the mud 
accumulation, but distal to the stomach the gut 
becomes so distended and the whole body 
inside the test becomes so compressed by it 
that it is difficult to imagine normal feeding 
and respiratory functions proceeding. Some 
of the mud must be lost through the arws anil 


ASCIDIANS OF SOUTH AUSTRALIA 


Ek 


Fig. 36. Curella eumyota. (Hallett Cove, 25 m). Individual removed from test. 

Figs, 37, 38. Ascidia sydneyensis. Fig, 37.—Individual from Tapley Shoal, 13 m. Fig. 38.—Individual 
from Wright I., 10 m. 

Figs, 39, 40. Ascidia gemmata, (Upper St. Vincent Gulf. 10-12 m). Fig. 39. -Individual removed 
from test. Fig. 40.—Diagrammatic section through branchial papillae, 

Figs. 41-43. Ascidia thompsoni. Fig. 41.—Dorsal lamina. Fig. 42.—Individual removed from test 
(Carickalinga Head, 5-6 m), Fig. 43.—Individual in test (aff West Beach, 8 m). 


Figs. 44, 45, Ascidia aclara. (Off Seacliff, 16 m). Fig. 44.—Whole individual, Fig. 45 —Individual 
removed from. test. 


6 PATRICIA KOTT 


uifial opening and until ubseryations are made 
on living specimens. it must be asstimeéd that 
the property of the distal part of the gut ta dis- 
tend ilsell in this way is characteristic af the 
species and results in the acctimulutions of gut 


contents al i greater rate than they are 
removed from the body. 
Ascidia gemmata Sluiter, 1895: 177. Kou, 


1966; 296 and synanymy. “Cokioka, 1967- 
149, 

New Recordy: Upper St. Vincent Gulf, off 
Port Gawler. off Glenelg. Previous Records: 
W. Aust. (Cape Jaubert to Albany)]—Hart- 
meyer 1919: Michaelsen & Hartmeyer 1928: 
Koil 1952, Vic. (Port Phillip Bay}—Koall 
1952; Millar 1966. N.S.W, (Port Jackson, 
Acruwarra)—Herdman 1899; Kott 1952. 
Qid. (Hervey Bay)—Kort 1966.  [ndo- 
Pacific (Indonesia) —Sluiter 1904; Tokioka 
1952; (Palao Is., New Caledonia, Matianas 
Is,, Caroline Is. Wake ts.) —Tokioka 1950, 

9G. TYaT. 

FIGS. 39. 40 

Meveriptions Externally the test is fairly thin 
and flaccid and is slightly irregular, The bran- 
chial aperture is terminal on a short cylindrical 
siphon, ‘The atrial aperture is on a similar 
but venecully shorter siphon Tron) the antero- 
dorsal aspect of the body, Both siphons are 
regularly prooved externally wlong their length, 
Individuals are attached by almast the whule of 
the left side. Internally the atrial siphon arises 
from half way dewn the body and is especially 
jong. The branchial siphun is alsa long inter- 
nally, There are circular and longitudinal 
museles around both the siphons and these 
extend only a Short distanee pusterior to the 
siphonk. on the left side of the body where 
there is nu musculature, On the right side of 
the holy the lonyitudinal qiuseles from the 
siphons mingle with the irregular meshwork of 
muscles which occupy the whole body wall on 
the right side. There is only a very narrow 
prebranchial area terminated anteriorly by very 
numerous branchial tentacles, and covered with 
minute papillae, The dorsal tubercle is 
fairly large circular cushion with a U-shaped 
slit turned to the fight and with the pos- 
tering horn turned in, The pentubercular 
ater is shallow and is completely filled by 
Vhe dorsal tubercle. The dorsal lamina is 
a broad, single membrane, strongly ribbed 
on both sides. The ribs of the dorsal 
lamina extend into pointed languets on the 
free margin, There iy a long ocural gland 
almost one-third of the body distant Fran the 


dorsal tubercle. The branchial sac is simply 
folded between each fongitudiqul vessel and 
has 4 to & stigmata in cach mesh. There are 
large spatulate papillae at the junctions of the 
longitudinal and transverse vessel and these ure 
expanded into rounded expansions on cither 
side of thelr base, The gut forms a Ueep 
double loop enclosing the gonads jn the pri- 
mary loop. The pole of the gut loop in the 
large specimens available in this collection does 
not catend anierior to the buse of the atrial 
siphon und is level with the anus, There ts. 
however, some variation accerding to the size 
of the specimens and jn smaller specimens 
(Michaelsen & Harimeyer 1928: Millar 1966) 
the gut loop extends anterior to the atrial 
siphon und occupies u relatively larger portion 
of the left side 


Remarks: This species hus been recorded often 
from) locations around Australia extending 
north 19 Indonesia and into the Pacific 
(Tukioka. 1967). The species is distinguished 
by the absence of intermediate papillac in the 
branchial sac. hy the heavily ribbed broad dor- 
sal lamina, and by the origin of the atrial 
siptraw from the middle of the body. Although 
in the present specimens the atrial siphon as 
long and directed anteriorly, in specimens pre- 
viously deseribed there js a great variation 
hoth in the length uf the atnul siphon and in its 
orientation ¢Michaelsen & Hartmeyer 1928)- 
Specimens have been deseribed with sessile ex- 
ternal apertures umd it is probable that the pre- 
sent specimens with short grooved cylinders 
represent more mature individuals. Exrernully 
the species resembles bouh 4. svdieyensiv and 
A. thompyoné and it is probable that in all 
these species the test is firmer and relatively 
thicker and the extemal siphons less evident 
in the younger specimens, while in older speci- 
mens the test becomes rougher externally and 
less transparent, and the external siphons dJeve- 
lop as short grooved cylinders. The body mus- 
culgture. concentrated on the right and on the 
siphons, is so arranged that the left side, fixed 
io the substrate, does not contract over the 
voluminous gut. In these specics the gut eceu- 
pies a relatively smaller proportion of the body 
wall as the individual inereases in size. In A. 
genimatad growth appears to increase the pro- 
portion of the body anterior to the gut, and 
although the point of origin of the atrial 
siphon remains about onc half ta two thirds 
of the distance down the body, the gut does 
not appear DO increase in sive gt the same rate 
as the rest of the body. The oricntation of 


ASCIDIANS OF SOUTH AUSTRALIA ar 


the gectum and the curvature of the gut loop is 
therefore reduced us growth proceeds. 1 is 
also possible that this cifferential growth caitses 
the varigtions that have been observed in the 
length and orientation of the atrial siphon, 
although this may also be affected by the oricn- 
tation of the body on the substrite, 


Axeidia muilacca ansiteliensis Hartmever, 
1928, resembles the present species in the pre- 
senge of a broad ribbed dorsal laming with the 
Iree margin produced into pointed projections 
sorresponding to the ribs, However. the spe- 
Ges is distinguished by the specially long exter- 
nal siphons. by the dorsal ganglion which js 
only one-ninth to ofe-thirleenth of the body 
length from thé dorsal tubercle, and by the 
amall stumpy cone-like branchial papillae as 
opposed to the sputulate papillae of 4. ge 
mao. Harimeyer's subspecies was recorded 
from a seasonally hvackish environment in 
Freshwater Bay, a considerable distance up the 
Swan River estuary from Fremantle Harbour 
and he regurded jt as an isolated endemic spe- 
cies, 


Ascidia thompsoni Kott. 1952: 312. 


Now Reeords: OF West Beach. Hullett Cove, 
Calickalinga Head, Previews Recordy: Tas. 
(Greai Taylor Bayi—-Kott 1952. 


FIGS. 41-45 


Pescripion: In smaller specimens the test is 
firm and almost glassy and transparent. Ante- 
rioly, expanded terminal ampullac of the test 
vessels are clearly visible through the test. Indi- 
viduals from 2 to 7 cm Jong are available tn 
the present collection. Both apertures are 
sessile. the branchial aperture terminal and the 
ttrial aperture two-thirds of the distance 
down the dorsul surface. Most individuals 
are firmly fixed by the whole of the left side, 
however the apecimen fron} Carickalinga Head 
is fixed posteriorly. The body musculature is 
present only on the right side, consisting af a 
mesh of dvansVerse and longitudinal vessels 
Internully the atrial aperture is on a siphon of 
variable length ising apposite, anterior or pos- 
terior 10 the external opening. The atrial 
siphon shows the same vanations in length and 
orientation as have been described previously 
for A. geminata (Michaelson & Hartmeyer 
1928; Millar 1966). Both siphons are well 
equipped with circular and longitudinal 
muscles, There arc about 40 branchial ten- 
tacles, a papillated prebranchial area, a shallow 
perituberculir area completely filled by the 
dorsal tubercle which generally has a sirnple 


U-shaped opening. In an especially lurge and 
Opaque specimen from West Beach (at % 1) 
there is a second opening to the feht ol the 
larger U-shaped opening. The dorsal ganglion 
is about half the bouy distant from the dursal 
tubercle, The dorsal Jamina is 2 wide mem- 
brane, dowble for about one-sixth of its length, 
The right section of the double membrane is 
plain, the left section is ribbed un the left. For 
the remainder of its fength the dorsal lamina 
is a single membrane ribbed on the left side, 
although these cibs do not extend to the outer 
margin of the membrane. There are minute 
ind irregular papilla-like expansions from the 
free border of the membrane tn its posterior 
extenl. Intermediate branchial papillae are 
generally present, especially in the posterior 
part of the branchial sac, The intermediare 
hranehial papillae are half the size of the pri- 
mury papillae, and bowh are pointed. ‘The gut 
is voluminous and forms a deep double loop 
which varies slightly in relation to the atrial 
siphon as the individual grows, us in 4. gen)- 
freilet, 


Remarks: Ihe dowble dorsal famina with 
slightly irregular membranous border pos- 
teriorly and the form of the intermediate and 
primary branchial papillae distinguish this spe- 
vies from the very similar A. genceta with 
whith tty geographic runge overlaps, The origin 
ind the variable orientation of the atrial siphon 
are shured by the two species, and in bath, 
owing to differential wrowth of the body, the 
gut loop is confined to the posterior half of the 
left side in latger specimens, Tt is of consider- 
able interest that the present species has been 
recorded only from fairly sheltered constal 
environments (subject however ta some wave 
action) in the present collections. while 4, 
geninlase was taken only from Offshore Benthic 
locations subject to currents in middle and 
\pper St. Vincent Gult 


Ascidia aclara Kot), 1952: 309. Millar, 1963: 
721; 
New Record: Off Seacliff. Previous 
Records: Vic, (Lakes Entrance, Port Phillip 
Bay)—Kott 1952, Millar L963, Cd, 
(Moreton Bay)—unpublished recurds, 
PIGS. 44, 45 


Description, There are two specimens in the 
present collection, maximum length 17 em and 
10 cm high The body is slightly darso-ven- 
irally flattened. The test is rigid anu encrusted 
with suind and shell particles and is produced 
into two tigid cylindrical tubes from around 


as PALRICIA KOTT 


the branchial and atrial apertures at the anterior 
end of the dorsal surface and from about onc- 
third of the distance ulong the dorsal surface 
respectively, The apertures afte completely 
sessile und lie at the base of these tubes, The 
hody nvseulature, within this rigid test, is 
reduced to strong bands ucross the dorsal sur- 
face posterior to the atrial aperture ind 
between the atrial and branchial apertures. 
Internally the specimens are exactly as pre- 
viously described with the branchial sue tormm- 
ing a fold across the dorsal tubercle. The gut 
forms ihe usual simple open Joop, opening 
adjacent to the alrtal upercure. 


Remarks; This unusual species apprars to be 
highly specialised for an existence on a sandy 
bottom, with the rigid tubes extending vertically 
from the apertures forming « permancnily open 
channel thraugh the layer of sand jo which 
the species is probably buried. It is probuble 
that the immediate environment outside the 
apertures is madified by these permanently 
open chambers to facilitate a less interrupted 
feeding process and confer distinct »dvantuges 
In locations where steady flowing currents and 
absence of sedimentation pertain, The species 
is also of considerable interest in that ils 
records are confined to the semi-enclosed 
waters indicated above. IL is possible that 
there Is a Wider, more continuous distribution 
on the continental shelf or, alternatively. that it 
represeuts a Telict population of a species which 
once hud such a continuous distribution on the 
open coast. 


Suborder STOLIDOBRANCHIA 
Family STYELIDAE 
Subfamily PaLyzornak 


Stoloniiy australis Michuelsen, 
Michaelsen & HMartmeyer. 
Kott, 1952; 253, 


New Records: Vipara Reef. Por Noarlunga. 
Previows Records: W. Aust. LAlbuny)— 
Michaelsen 1927; Michaclsen & Hartmeyer 
{4928 ‘Tas, {Spring Bay}—Kott 1952, 


FIG, 46 


Dexcriptlon: Rounded, sandy. stalked or sessile 
dividuals connected to basal stolons, 0,6 to 
(7 em maximum diameter. The colonies in 
the present collection are encrusting specimens 
of Pyne irregularis and Polyearpa peditnen- 
love, The apertures are both sessile on the 
upper surface. There ate two folds on either 
side of the branchial sac with 6 to 9 internal 
longitidinal vessels. The gut loop is simple 


1927; 
928: 


202, 
352. 


and open with a gastro-intestinul ligument 
enclosing a rounded endogurp in the pole, The 
short stomach has about 18 Folds. tt is. reduced 
jn diameter at either end and his a thick pyloric 
caecum of moderate length. Gonads are not 
malure in the present specimens und their 
arrangement could not be determined. The 
ligaments anchoring the gut to the body wall! 
extend in a tow along the lateral aspect of the 
intestine. There are also large hygaments 
anchoring the stomach and the pole of the gul 
loop. 

Remarks: This species appeyrs to he confined 
to the southern coast of Australia, but has been 
recorded only from locdions away from the 
open coast, It is inconspicuous. however, and 
iL is possible that its occurrence in protected 
locations on the open coast has been over- 
looked. [nthe absence of mature gonads the 
species miy be distinguished from dmplicarpa 
dipiycha by the low rounded branchial folds, 
the presence of a curved pyloric caecum and 
the less developed musculature. 


Stolonica carnosa Millar, 1963: 734. 


New Record: ‘Vipara Reef. Previnses 
Record; Wo Aust. (Cottesloe), 
FIG. 47 


Description: The colony is oval, 3 em long, 2 
em wide and | cm thick and, as in the type 
specimen, hus developed around iin algal stem. 
The 4-lobed apertures of zooids are close 
Togelher an slight swellings all yround the outer 
surface which is encrusted with sand, There 
is nO sand inside the colony, Each indivatual 
ix Jorsa-ventrally flattened and most of ils left 
side is directed toward the centre of Lhe colony. 
There ure 2 folds on euch side of the branchial 
sur with internal longitudinal vessels according 
to the following formula. E O( 434491 DL, 
There are only 3 stigmata between the endo- 
style aod the ventral fold. 

The cut forms a rounded loop and the rec- 
tum turns anteriorly and dersally at a sharp 
angle. The stomach is pyriform, narrowest at 
the curiae end, has 15 pacrow folds and a 
very long, curved pyloric caecum in the pole 
of the put loop. There is a gastro-intestinal 
ligament and ligaments connecting the gul loop 
to the body wall as in Distemuty diptyche (see 
Kort 1952). The anus is 2-ipped. The gonads 
are in single rows on cach side of the endo- 
style, The testes arc fMask-shaped and the 
ovaries contain 3 exes of varying sizes, and a 
testis and an ovary are generally loosely asso- 
ciated so that there are 6 to 7 hermaphrodite 
gonads on each side af the body, 


ASCIDIANS OF SOUTH ATISTRALIA ay 


Remarks; Although in Millur's specimen the 
testes and Ovaries appeared often to be sepu- 
rate, the condition and the artungement of the 
gonads in the present colony suggest that this 
is Mole apparent than real. and may depend 
on the relative stages of Uevelopment af the 
ovary 


In Millar’s specimen the stomach is folded 
internally but externally the tokds were pro- 
hubly obscured by the membrane covering 
them. The course af the reetum in the present 
specrmen also differs from Millar's specimen 
and is bent back against the gut loop, prohahly 
by dorso-ventral flattening of the individual. 
The extent of vhis dorso-ventral flattening. 
therefore, is am individual, rather than # speci- 
fic. character. 


Oculinaria australis Gray, (868: 564. Katt, 
1952: 25) and synonymy. Millar, 1963: 
73d; 1956; 369, 

New Records: West 1. (Scal Rock), Wright 
I. Previous Recérdy: W, Aust. (Fremantle 
to Albany )—Gray 1868; Michaelsen & Hart- 
meyer 1928; Kott 1952; Millar 1963, Vie. 
(Port Phillip Bay)—Millar 1966. 


Description: Colonies of the usual form with 
numerous zuoids closely coalesced, identified 
only by the paired apertures on wart-like 
siphons from the anterior surface of each 
zanid which project slightly from the other- 
wise campact colony. The test is very brittle 
and completely (Mpregnaled with sand. There 
are 4 branchtal tolds on each side of the body 
with 4+ to 8 longitudinal vessels on cach Fold 
and about 4 between the folds, The gut loop 
is as previously described, with about [8 spiral 
folds in the stomach wall. No pyloric caecum 
has heen detected. There is an elongute gas- 
tric gland reservoir extending between the 
stomach and the intestine, There are up to 9 
lonz gonads on the right side of the hady, a 
lurger number than hus previously been 
recorded for this species. There is a single 
row of testis lobes beneath cuch short ovary. 


Remarks: The species is well adapted, by its 
compacted form, far the occupation of turhu- 
lent locations and, in fact, it has been recorded 
only Erom the exposed open coast, Externally 
iL resembles colontes of Palyandrocarpa spp. 
from which it is readily distinguished not only 
hy the location of the gonads on one side of 
the bady, but also by the spiral course of the 
stomach folds, the presence of 4 gastro-jntes- 
tinal reservoir and the form of tbe gut loop 


Subfamily moveyoirivar 


Botrylnides leachl (Savigny), Michaelsen & 
Huitmeyer, 1928: 341 and synonymy 
Millar, 1952: 24.1962: 177) Kort, | 952) 
233; 1966: 297 
Boiryllus leachii Savigny, 18107: 7. 

New Records: Tipara Reef, Port Noarlunga, 
West 1, Wright l. Previews Recarey: W. 
Aust. (Geraldton to Albany) —Michaelsen & 
Hartmeyer 1928; Kott, 1952, N.S.W. (Port 
Jackson)—Herdman 1899. Qld. (Moreton 
Bay}—Kott 1952; (Sarina)—unpublished 
record, Northern Territory ¢Darwinj— 
Kott 1946, New Zealand (Hauraki Gulf )— 
Michaelsen 1921; Brewin 1948; (Stewart |.) 
—Michaelsen 192[; (French Pass)—-Sluiter 
1900; (Otago Harbourj—Brewin 1946; 
{Auckland 1,}—Bovien 1922. South Africa 
—Hartmeyer 1912; Millar 1962. The spe 
cies 2s alsa Known from the North Atlantic. 
the North Sea and the Mediterranean and 
Adriatic (see Hartmever 1923, Arnbiick 
1923, and Millar L952). 


Deseription; Living colonies from Ocdipus 
Point, West T. have u colourless matrix and red 
zooids, while in those from Port Noarlunga the 
mulrix (s transparent and the zooids yellow- 
bright orange. All the colonies have trans- 
lucent test and purple zooids in preservative 
Colonies form fMattencd, long, Iohes with a 
shore stalk. There ate circular to oval svsiems 
of closely packed zoids. The test i< firm and 
transparent. The system af zooids are arranged 
in rows along the Jength of the head, These 
systems May appear to be confluent and form 
almost continuous rows, but in fact separate 
cloacal openings remain in the centre of a 
limited number of zooids and dis¢rete circular 
to oval systems are niainivined) There are 9 
lo 12 rows of about 20 stigmata. “Phe stamach 
is long, with 10 folds and u very short caecum, 


Remarks: The form of the colonies és very 
similar to those of B, magnicoecuim bul the 
circular systems and firm test, with common 
cloacal openings along the sides of the lobes 
ire distinctive. The shape of the stomach and 
the form and length of the pyloric caecum is 
similar to the condition found in B, nigrum. 
However, the smaller number of rows of stig- 
Mata with more stigmata in cach row also dis- 
tinguishes this specics from both B. magai- 
coecum and from 8. nigrum. Records for this 
species extend from the North Atlantic to the 
Medilterrancan and Pucific Oceans, and from 
All around Australia. It is not known from the 


30 PATRICIA KOTT 


Indiun Ocean beyon the West Australian coast 
nor is it known fram the South Aflantic. 


Botrylloides nigrum Herdman, 1386: 50, Van 


Name, 1845: 227 and synonymy. Katt, 
1952: 257 
Surcvlniviiaides  jacksonianam  EHerdman, 


1899: 102. 

penbeyestatat pamasniny Herdman, 1899; 
New Records’ Port Gawler, elf West Beach- 
Off Seneliff, Curickalinga Head, Rapid Head, 
West 1, (near Penguin Rock, Seal Rock). 
Wright | Previous Recards; W, Aust., 5. 
Aust. Vie—Kott 1952. N.S.W. (Port Jack- 
som t— dlerdmarn 1899; Kott 1952. Old— 
Kote 1952) Indo-Malaya (Ceylon)—Herd- 
man 1906: (Red Sea)—-Michaelsen 1919. 
Fast Africa-—Shuiter 189%, Michaelsen 1918. 
South Atrica—Hartmeyer 1912. The spe- 
cies is also recorded from the Curibbean 
region (Van Name 1945), 


Deyeription: Colonies investing sheets some- 
times extended into irregular lobes, The zooids 
wie arranged in long double row systems well 
separaled from one another with transparent 
test between In preservative the zooids are 
purple-lback with the pigment contained in 
cells in the hady wall. The colour of the pre- 
served specimens does not reflect the variations 
in colour of the hving specimens which irc: 
“dark blue and bright purple” zaoids {West §.): 
ay “vellow and musturd” (off West Beach), 
There ave 16 rows of about 12 stigmata with 3 
internal longitudinal vessels of each side of the 
hranechial sac. The atrial opening exposes the 
anterior half of the dorsi! surface of the bran- 
chial sac, but the lip from the anterior border 
of this opening ts not especially pronounced. 
Ihe stomach is the usual long organ charac- 
teristic wl this species, with 10 folds, I is 
wider at the cardiac end and reduced in width 
at the pyloric end where there ty a very short 
cuecuin. 


Retnarks; Aithough the variation in colour and 
the irregularity of the colunies make this spe- 
cies difficult io identify in the field, the: shape 
of the stomach with its shurt caccum and the 
widely spaced double rows of zooids are dis- 
linctive. [ts recorded distribulion is wide in 
the Indian Qceun and from the West Indies. 
At this stage there is no known character avail- 
uble (o indieute that all these records refer to 
more than the ane species with an almost cir- 
cumpoler distribution. in the southern temper- 
ate region, absent onty from the middle and 
eastern Pacific Ocean. 


Rotryloides magnicwecum Huttmeyer. Kott, 
1952: 258, Millyr, 1966: 368. 
Borylloides nigrum var, magnicoecum Hart- 
meyer, 1912; 271, 

Reavllus magnicoecus. Michaelsen, 1923b: 
50; 1923: 4, Mivhaclsen & Hurimeyer, 1928; 
331 and synonymy. Hustings, J93L. 79. 
Brewin, 1951: (09. Millar. 1958: [95> |9n2- 
175. Tokioka, 1967; 153, 

Bolryllay auceps Michaelsen & Harimeyer, 
1978; 335. Millar, (963: 774. 

Palvexelas rufus Oka. 1927) 6b8 

Betevilius refs. Tokioks. 1943h: 240 


New Records: Off West Beach. West b. 
Wright J. Previews Reecards; W. Aust 
(Shark Bayj—Michwelsen & Hartmeyer 
1928. S. Aust, Tas.—Kott 1952, Vic 
(Port Phillip Bay}—Millar 1963, 1966 
N.S.W. (Port Jackson) -Herdman 1891; 


Millar 1963. Qld. (Great Barricr Recf}— 
Hastinus 1991. New Zealand (North b.j)— 
Michaelsen 1921: Brewin 195). Japan— 
Tokinka 1952; Oka 1927. Chinu (Hony 
Kong)}—Michaclsen 19232, Tokioka 1967, 
tadian = Ovean (Paumbur) — Michuelsen 
19235. South Africa—Hartmeyer 1912; 
Millar 1955, 1962. South West Africu— 
Hartmeyer 1913; Michaelsen (915. Natal— 
Michnelsen 1918, 1921. Rurope (Portugal) 
Tyar—Michaelsen 1923b; (Mediterranean) 
‘var. Michaelsen 1923b. 


Meserprons The living colonics from West 1. 
are “bright yellow" although other specimens 
are “oreyish with pale zoojds”, In preservative, 
however, all the colonies are purple owing to 
the pigmentatiow of the zovids which shows 
through the very solt transparent test. The 
colonies in this collection always consist of 
soft, long, narrow, Mattencd, stalked lobes with 
zooids arranged in closcly set double rews 
running parallel to the Jength of the lobes 
Zooids are absent from the stalks. In preserved 
Specimens there is always an accumulation of 


dark pigment at the top of the endostyle and 


on either side of the bine of the branchial aper- 
ture Common cloacal openings are ulwiys 
present around the free end of the lobe as. in 
Syvrezoqd spp. 

There ure |4 rows of stigmata in the present 
specimens with 3 tw 4 stiymata between the 
Jongitudinal yessels. The stomach is short and 
rounded with 9 folds und a long caccum curv- 
ing into the pole of the gut loop. 


Remarks: Millar (1963) regards the form of 
ihe colony of the Australian specimens (long 
stalked lobes) as providing & character which 
distinguishes it from the South African forms 
which are irregularly lobed and tnvesting, as 


ASCIDLANS OF SQUTH AUSTRALIA 3) 


are Brewin’s specimens from New Zealand, The 
closely set double row branching systems are 
present in all the specimens represented in the 
synonymy above and all these specimens have 
the characteristic shart, rounded, stomach with 
a long curved caecum. distinzuishing them 
From other spectes of the genus It is possible 
that the Australiin members of this species 
may represent a geographic subspecies charac- 
terised by the particular form of the colony 
with terminal cloacal apertures and close-set 
double rows of zooids parallel to the longi- 
tudinal axis af the head. Bostrylinides leacht 
colonics are similurly lobed but the cloacal 
apertures are present along the side of the head 
hetween the double raw ol zoeids- 


Botryllus schlosseri (Pallas). Van Name, 1945; 
220 and synonymy, Kott. 1953: 259 
{purt), 

Aleyaniumn yehlasseri Pallas. 17662 454. 

Non Aeeryllus sehlosserd. Kott, 1952. from 

Hamelin Bay and Green Pools, W.A, 
New Recera: OF Hallett Cove,  Previnne 
Records: W. Aust. (Shark Bay, Fremantle}— 
Hartmeyer & Michaelsen 1928: Kou 1952 
Vic, (Port Phillip Bay)—Millir 1966. Else- 
where the species has a wide distyibution 
from the Faeroe Is. and southern Norway. 
the British Istes, the North Sea. the Mediter- 
fanedn, Adrintic and Black Seu; from the 
eastern and western seaboards of the U.S.A- 
and from New Zealand (see Van Name 
1945), 


The locul abtindance of this species and its 
occurrence on wharf piles. ship hulls, buoys, 
ete. in shallow water has been pointed out by 
Van Name (1945). ‘This wide cosmopolitan 
distribution suggests that, like Ciona intosrin- 
«lis, the species favours sheltered locations and 
is transported largely by ships. 


Description. The specimens are delicate and 
invest the sea vrass Poyidania australis, The 
lest is almost completely transparent and the 
zoaids are pale grey, Zooids form srnall cir- 
euwlar systems which ate crowded close to- 
gether in the test. The zooids are relatively 
short. with only about 8 rows of stigmata, The 
atrial aperture is on a siphon produced to a 
varying extent and the upper margin of the 
aperture is produced into a lip. There is a 
conspicuous pyloric caecum: with a large bulb- 
like expansion on its free end. The stomuch 
has about 10 very fine folds. is longer than 
wide, and is only of slighty greater diameter 
than the fest of the gut. Developing embryos 
are present in the peribranchial cavity of some 


of the zovids, bul on the right side of the hos 
only. 

Remarks: The zooids ina colony of the present 
specimens are identical with those described 
for Borrvilix gracilis Harimever & Michaclsen 
1928; Millar, 1966, from Shark Bay, Western 
Australia and fram Port Phillip Bay. Millar 
(1966) regards this type of thin transparent 
colony us a species distinct from B. sechlosseri. 
Juvenile colomes of B. schlosseri as described 
by Verrill (Verrill & Smith 1873) are identical 
with the presont coluny and the zooids are iden- 
tical with those previously described for this 
species especially in regard to the atrial open- 
ing, stamach anc pyloric caecum. and it Is 
unlikely that  graciliv is distinct from &. 
schlosseri. 


Subfamily stveciwae 
Cnemidocarpa etherldgii (Herdman) 
Stycle etheridgii Herdman, 1899; 38, Kolt. 
1952: 219 and synonymy; (964: 129 (F. per- 
sonata), Millar, 1966: 370, 
New Records: Tapley Shoal. off West Beach, 
West J. (off Oedipus Point), Wright . Pre- 
vious Records: W, Aust. (Vriga 1). 8. Aust. 
(Spencer Gulf and St. Vincent Gulf}. ie. 
(Phillip 1,1 —Kott 1952: (Port Phiflip Bay) 
—Millar 1966. Tas. (D’Entrecasteaus 
Channel}—Koatt 1952, N.S.W. (Part Juck- 
son, Port Stephens)—Herdman 1899. Qlu- 
(Morcton Bay)-—Kott 1964. 'Uhe species is 
known intertidally and down to 30 m. It 
is abundant in St. Vincent Gulf on sanity 
boltoms at 7-20 in with slow currents, and 
On Open coasts in decper water of 20-30 m 
(Shepherd, pers. comm.). 
FIGS. 48, 49 
Description: Individuals are large. up to 11 em 
high, rounded and of greatest diameter pus- 
lertorly. gradually reducing in diameter ta the 
terminal branchial aperture. The terminal 
branchial aperture is sometimes curved. The 
atrial aperture ts on a slight rounded projec- 
tion from about half way along the dorsal 
surface. Colour of living specimens Varies Fram 
pale cream to bright vellow (most often the 
fatter), In preservative the lest is white and 
Opaque, with longitudinal furrows converging 
to the branchial aperture on that part of the 
body anterior to the atrial aperture, The test 
is thin and leathery, There are up to 25 inter- 
Nal longitudinal vessels on the folds and up to 
7 between, although in some specimens there 
are as few ys 4 internal longitudinal vessels 
between the folds. There are 6 stigmata per 
mesh. 


42 PATRICIA ROTT 


Fig. 4G, 
Fig. 47. 


Individuul bisected along the ventral surface, 


and endocarps in body wall. 
Fig, 50. 


Polyearpa clavari. Whole individual, 


Stalonicu australis, (Tipara Reet), Gut loop. 

Stolonica carnosa. (Uiparn Reef). Right side of body removed to show organs on left 
hody wall. 
Figs. 48, 49. Cnemidocurpa ctheridgii. (Tapley Shoal, 13m). Fig. 48—Whole individual. Fig, 49.—- 


branchial suc removed, showing gonuds 


Figs. 51, 52. Polvearpa pupillata. (Of Glenelg, 15m). Fig. 51,—Body wall on left showing gut loop, 
gonads and endocarps. Tig. $2.—Individual showing gonads on right side of the body. 

Figs, 53-36. Polyearpa pedunculata, Fig, 53.—Individual from Aldinga (10-25 m). Fig, 54—indivi- 
dual from Taptey Shoal (24 m). Fig. §5.—Individual from West I. (25 m). Fig. 56—: 


Gat Joop and endocarp, 


The gut forms a gently curved, fairly narrow 
loop across the left side of the posterior end 
of the body, enclosing a Jong narrow curved 
endocarp which is continuous with the body 
will on both the right and left side. The gut 
loop is almost entirely posterior to the bran- 


chial sac: the elongate stomuch und proximal 
part of the intestine forming the proximal limb 
of the gut loop lie almost in the mid line pos- 
tero-ventrally and the distal limb of the gut 
loop passes to the Teft of the posterior end of 
the branchial sac. The gut loop is almost 


ASCIDIANS OF SOLTH AUSTRALIA 33 


entirely embedded [hn the thickened body wall 
und i covered by endocatp which encloses. the 
left gonad (in the curve of the gut) and extends 
ventrally iacross the pole of the gut loop to 
join the thickened body wall ventrally and pos- 
terjorly, The pole af the gut loop thus projects 
into jt pocket in the thickened body wall, The 
ovsophagus is short and the stomach is long 
and elliptical with internal longitudinal glundu- 
lar folds, The anal opening has a smooth 
border, 


There ure one or two flask-shaped gonads on 
the myht side of the body. On the lelt the 
gonad is embedded in a single large cndocarpal 
thickening Of the body wall, Here there may 
he a single branched gonad with single 3 and ¢ 
ducts emerging from the endocurp and directed 
to the utrial aperture “This condition may 
have resulted from the fusion of two gonads. 
In another specimen there are lwo discrete 
yonuds embedded in the teft side of the body 
with their own sets of cf and © ducts emerging 
from the endocarp. The testis lobes are 
enclosed by the ovarian tube as is characteristic 
of this genus. 


Remarks: The present farge specimens con- 
form with those ascribed (Row 1952) to the 
“erheridgii” condition af this species, It is 
most probable that this distinction relates only 
lo the stage oF maturay of the individual, where 
the “persuran” condition represents Jess 
mature individuals. Both forms hive been 
taken [rom the same locations in both cast and 
western Australia. 


Polycarpa clavata Hartimeyer. Millar. 1963: 
723. 
Polvearpa mau (Quoy & Gaimardy f. 


chwaty Hartmeyer, 1919: 40, Michachen & 

Hartmeyer, 1928: 363. Kott, (952: 23h, 

Tokiokn. 1961; 123, Vasseur, 1967; 133. 
New Revoards: “Vapley Shoal, near Marion 
Light, off Troubridge Light. Previons 
Reoordys: W. Aust, {Bathurst J, to Rottnest 
[y—Hartmeyer 1919; Kort 1952: Millar 
1963. Pucific (Noumes, New Ciledonia) 
Tolkinka 1961: Vasseur 1967, 


FIG, 50 


Deserfptlon: Large stalked specimens [rom 
fawn to reddish-brown The test is very soft 
and gelatinous and the surface is marked with 
rounded longitudinal ridges which are some- 
times interrupted horizontally. The branchial 
uperture is on a short siphon from the basal 
une third of the dorsal surface. <lirected to- 


ward the substrate. The atrial aperture 7s 
sessile and inconspicuous from the middle third 
of the dorsal surface. The upper, or postenor. 
end of the head is high and rounded. The 
stalk. about the same leneth as the head. is also 
thick und fleshy, wider toward the base in the 
larger specimen, and bulbous, or. in smaller 
specimens. (Marion Light) fairly narrow In 
the smaller speciniens there ure randomly dis- 
tribuled concavilies, surrounded by well defined 
lips, on the sides and base of the stalk. These 
concavilies are richly supphed with blaud ves- 
sels which end in terminal ampillac in the base 
und lips of the concavily, Ft is possible thul 
these organs are involved in the fixing of these 
individuals to the substriite, especially is they 
do not appear to be present in the harger speci- 
mew where the surface test of the stalk is wni- 
fermly transversely ridged. 


The musculature is rather diffuse in the thick 
body wall which is produced into a tongue-like 
projection extending about ong third of the 
distance down into the stalk. ‘The stulk ix cam. 
posed of solid test material for the remainder 
of its fength. The dorsal tuberele is large, com- 
pletely filling thy peritubercular area und has 
complicated, convoluted and interrupted oper. 
ing, There are 4 branchigl folds on either side 
of the body, samctimes only apparent as jin 
accumulation of longitudinal vessels, The 
branchial sae does not project into the anterior 
tongue of the bady wall where W projects inte 
the stalk, 


The gut forms a double loop confined to the 
posterior part of the body. ‘The anal horcer 
has small rounded lobes. Endocarps enclosed 
in Ue gut Jonp may he subdivided terminally 
into two or more branches. Gonads, more or 
less in 3 rows down each side of the body wall. 
are “fool” shaped. fixed to the body wall bv 
the metaphorical “ankle”. with the “toe” point- 
ing Joward the atrial aperture, 

There we numerous upright endocarps scut- 
tered over the body wall between the gonuds 


Remarks; The dorsal tubercle of Polyearpa 
pedata Herdman (Sryela pedata) which Han- 
meyer ¢1919} listed as a synonym of the pre- 
sent species is distinguished by the presence of 
humerous pil-like openings while the dorsal 
tubercle of the present species. although com- 
plicated, has a convoluted slit-like opening 
interrupted several times along its length. The 
Present species appears closely related to 
Polyeurpa longifermis Tekioka (Kott 1966), 


34 PATRICIA KOTT 


which has similat gonads and uppeurs to be 
distinguished only by the orientation of the 
budy, the absence of the distinctive stalk and 
the simple opening of the neural gland. Paly- 
cirpea ottoteny Herdman (1399) has a similar 
convoluted opening en the dorsal tubercle, 
sometimes broken into several openings ulong 
its length. The gomads in P. arrelens, however. 
are upright. 

Millar (1963) drew attention to the ditter- 
ence between P. aurata (Quuy & Guimard) and 
the present species first described as P. arate 
chivaa Hartimeyer. 


Po data, Hastings, 1931, ts described as 
ugrecing, well “with Hartmeyer’s (1919) and 
Herdman’s (/. safeara; Berdman |886) des- 
criptions”. Hartmeyer’s deseription.. however, 
is Of Polcarpa aurata F elavara (<P. clavata) 
and it is With PL aurata ¢>P. suleata) thar 
Hiusting’s specimen is identical, A re-examina- 
(ion of the type specimen of P. atraie var. 
plana Herdnvan. 1899 from Port Jackson has 
shown that its gonads are also the usual short 
polycurps of ?. anrare which is now knoWwo 
from Port Juckson and the Great Barrier Reet 
und from the Indian Ocean. Mulaya, and Indo- 
nesia The ranye of P-avrete, therefore, dors 
now overlap that of P. efavera, 


Polycarpa papiliata (S|uiter). 


Stvela pupitlata Sluiter, 1886: 192. Tokika, 

1952: LI7. Vasseur, 1969: 925, 

Palycarpea intéstitata Kott, 1952: 23k. 
New Records: Tipara Reet, olf Port Gawler, 
uff Glenele. Aldinga “drop-off” Previous 
Records: NSW, (Port Jackson)—Kott 
1452. India) Oceut (Madagascar)— 
Vasscur 1969, Indonesia Sluiter 1886: 
(Arafura Sea}—Tokioka 1952. 

FIGS. 5], 52 


Descriprion: Small ugeregates of individuals, 
the posterjor fest sometimes extended into a 
short stalk, The branchial aperture is terminal, 
the alrial aperture one third t one half of the 
distance along the dorsal surface, Both uper- 
tures are sessile. The test ix tough, rough and 
wrinkled exterqully, with some sand and algae 
irregularly adhering but yenerully the surface 
test is muiked. The body musculature consists 
of a moderately thick continuous external coat 
of circular muscles with longitudinal bands 
internally. The dorsal tubercle is a large blister- 
like swelling with a simple U-shaped opening; 
i} completely fills the V of the peri-tubercular 
area. There are 4 wide overlapping folds with 
about 15 jaternal longitudinal vessels on ech 


fold and 3 wo 4 between folds. There are 4 
§ stigmata in cach mesh. Anteriorly the endo- 
stvle Follows a winding Goutse, which ix effected 
by the subdyvision of transverse vessels and 
multiplication af the puntber of rows of stiz- 
mata ventrally, in a localised region slong the 
anterior cxtent of the endostyle. The gut forms 
a hnrizantal loop in the posterior end of the 
body. The stemach is olliptical with Jongi- 
tudinal striations. The rectum extends ante- 
tiorly toward the atrial opening. The anul 
border is broken up into 14 long fingerstike 
lobes, Tall endecarps are present im the vut 
loop and seatiered over the body wall. Seven 
ta 12 aval to elongate polycarps pre present in 
1 to 2 rows in the centre of cach side of the 
body, directed lowand the atria] aperture. These 
polycarps ute fixed to the body wall wong their 
whole length Ino smaller specimens with 
smaller immature gonads there are more often 
2 rows of polycurps, anu. as the gonads increase 
in length and the body length increases, (hese 
rows upper to merge inte au single inreguilier 
row, while in a single specimen with well deve- 
loped gonads there is only a single regular row, 


Remerky: The present species resembles Paly- 
cerpa clavate (Hartmeyer), Po lavnet/urreis 
Tokioka and P. attellens Herman, in the tall 
endocarps enclosed ih the put loep, but (5 dis- 
tinguished by the rows of recumbent gonads 
fixed alone their whole length to the body wall. 
The anal lobes also resemble those of P. 
atollers and PQ lengiforpis, 


The form of the body, the position of the 
attial aperture, che form of the dorsal rubercle. 
and the form and arrangement of the gonads 
are similar to P. emrctinurdte (Sluitert, Vasseur, 
1967, which is distinguished by its short oval 
stomach. greater number of rows of gonads und 
greater number of internal longitudinal vessels 
between the branchial folds, 

Cnemidocarpa omiutedagascariensty — plelagas- 
carteusts Harumeyer from Madagascar and C, 
moadegascarienyis reguliy Michelsen from New 
Zealand (see Kott 197]a) also resemble the 
present Species in external appearance and in 
the arrangement of gonads, and are cistin- 
guished principally by the greater length of the 
gut loap and greater number of internal lonyi- 
tudinal vessels; between the branchial foldy 
The papillac on the branchial sac described by 
Sluier (1886) are not present cither m the 
South Australian specimens or tn the specimens 
fram the Arafura Sea (Tokioka 952). Tr ts 
possible that Sluiter mistook particles adhering 
to the branchial sac for papillae, 


ASCIDIANS OF SOUTH AUSTRALIA ss 


The species has a wide geographical cistribu- 
con from Indonesiq and apparently around the 
cast Goast of Australia, from focky subsirates 
I) sheltered localities, or in Olfshore Benthic 
locations where there are slight currents. 

Populations of Lhis species du not appear to 
be dense and tecards are few, 


Polycarpa pedunculita Heller, 1878; 106. Kott, 
1952; 237 and synonymy. Miliar, 1966: 


369, 

Falscurpa e@bsenra. Kott, (952: 245 und 
spnanynty, 

alycurpa steplienensis Herdmun, 1899; 45, 


Koit, 1952: 232. Millar. 1963, 726, 
Polyearpu moehii, Katt, (952: 244 and syno- 
nymy; 1966: 299. Vasseur, 1967: 136, 
Polycaurpu obrecta. Kot, (52: 242 (nat P. 
ohrecta Traustedt), 
New Records: Vipara Reef. Tapley Shoal, 
near Marion laght upper St. Vincent Gull, 
off Port Gawler. off Semaphore. off Grange, 
aff West Beuch, off Glencly, off Broadway, 
otf Hallett Cove. Port Noarlunga, Aldings, 
Cinckalinga Heal, Rupid Head. West L., 
Wright |. Previnns Records: W. Aust. (Cape 
Ivuhert to Bunhburvi—Hurtmeyer 1919; 
Michvelsen & Haortmeyer 1928; Katt 1952, 
S. Aust, (Reeveshy 1). Vie, ( Balnarring 
Beach )—Kute 1952; (Bass Straiti— Heller 
1878; Michaclsen 1905; (Port Phillip Bay) 
—Millur 1966. N.S.W. {Port Jackson, Two- 
fold Bay}—Herdman 1881. Qld, (Mereton 
Bay}—Kotl 1964, The species hus dlsu been 
recorded fram New Claledonia (Vasseur 
1967). 
FIGS, 53-56 
Description. This ts by tar the most common 
ascidian in St. Vincent Gulf and is very vari- 
able to external appeatance, The colour of 
living specimens from Port Nourlunga has been 
described us “bright to pale yellow”. ‘Vhese 
specimens are black to greenish in preservative. 
Most often living specimens are sandy with a 
“reddish linge” to “reddish brown" becoming 
brown to purplish brown when preserved in 
formalin. ‘They are slightly laterally flaiened 
and almost oval shaped, and are most often 3 
to 4 cm long and 2 to 3 cm wide, Larger 
specimens up to 8 cm long ure usually grecnish- 
black in preservitive. The apertures are ses- 
silé. the branchial aperture terminal but direc- 
ted slightly to the side, away from the dorsul 
surface, and the atrial aperture one-third of the 
distance down the dorsal surface, 
The test. is tirm and gelatinous and the sur- 
face is generally smnuch and naked, There is 
often. hawever, 4 light encrustation ol sand 


or the test may be more heavily encrusted. or 
may heeome almost brittle with included sand, 
In larger specimens the test becomes thinner. 
more flaccid and leathery, 


Posteriorly the dest may he produced into a 
narrow atalk up to half the length of the bouy, 
or the body may taper gradually from a straight 
upper or anterior surface whene the branchial 
aperture ts central and the atrialaperture is on 
the antero-dorsal corner. The posterior end of 
the body, with or without iw stalk, may be pro- 
duced into root-like structures, or the incivi- 
dual may be fixed to the substrate by the 
postero-ventral surface. 


Ihe body wall is Jight tu dark brown, 
hrownish-green. greenish-black, or black. It 
is not very closely adherent to the test and js 
thick, firm and very muscular with internul 
longitudinal bands and a continuwus. thick 
eXternal coat of circular muscles. Both layers 
of musculatiire are often embedded in fleshy 
non-muscwlar tissue and yvencrally spherical 
vesicles are embedded in the smuscle layers 
interrupting the regularity and cominwity of the 
fibres. The hudy wall is more flaccid |W larger 
specimens. 


There ate ubeut 100 simple tentacles at al 
least 4 orders. The prépharyngeal arca has 
small papiive and is of moderate width, The 
dorsal tubercle varies and is sometimes small, 
in the centre of a fairly Large peritubercular 
area. Wt is sometimes much Jarger but never 
completely fills the perituberculur area. The 
opening forms a U with borns turned in or out 
and directed to the side, antesiorly or poste 
riurly and in larger specimens may be inter- 
rupted. The dorsal lamina is a plain edged 
narrow membrane. The branchial folds are 
low and rounded with 2 ta 3 thick internal 
longitudinal vessels between the folds and LL to 
13 on the folds. There are f to B stigmara 
in each mesh between the folds but on the folds 
the jnternal longitudinal vessels are more 
crowded together. There are often vesicles, 
similar to those embedded in the body wall, 
embedded in the branchial vessels and in the 
dorsal tubercle, The gut is confined to the pos- 
terior cnd of the body distal to the atrial aper- 
ture. ‘The intestine forms a short rounded loop 
enclosing 4 circular endocarp. The stomach 
itself is elliptical with pronounced fulds, There 
may be a second small endocarp separating the 
rectum from the ocsophagus as the former 
extends anteriorly toward the base of the atrial 
opening. In smaller specimens the anal border 
is broken into 7 sometimes subdivided rounded 


Ais) PATRICIA KOTI 


lobes. In larger specimens there are up to 25 
lobes. The circular endocurp enclosed by the 


gut appears to be the miyjor mechanism anchor- 
ing: the gut loop to the badly wall and is con- 
fluent with the connective tissue surrounding 
the gut. There are 20 te 50 short oval poly- 
carps on the Jef and 25 to 60 on the right 
These are sometimes, but not always, embedded 
completely in the body will. Wher completely 
embedded only the openings of the ducts wre 
apparent us holes. in che inner surface of the 
hody wall. Primarily there appear to be about 
3 longitudinal rows of polycarps on euch side 
of the body. As each polycurp increases in 
length it subadivides and new gonoduols open 
from the proximal half to form secondary rows 
of gonads overlapping the primury row closest 
to the atrial opening. Wi is possible that. this 
process, resulting in Increases in the number of 
polcarps present, explains the great variacion 
in the number recorded for this species. 


Remerkss Mivhuelsen & Hartmeyer (1928) 
dew. attention to Lhe similarity between species 
listed in the s¥nonymy above and suggest that 
P. obyeura is a vuriely of PL peduneulata CP. 
viridis).  Michaelsen regarded = Polvcurpa 
moebu, however, as 4 distinct species charac- 
ierisedl by differences in the gut and gonads. 
In this collection there are individuals demon- 
strating every condition previously deseribed 
tor P. pedtinenlata, P. moebil, P. obseura and 
P. siephenensiy. There are specamens demon- 
vtrating every condition from stalked or ranted 
io sessile individuals; every colour und every 
condition of the test is found and there ps con- 
siderable yariation in the number of polycarps 
and the extent lo whieh they are enthedcled. 
The gut loop is always constant and encloses 
the cireulitr endocarp which has. a pointed tip 
dorsally, The thick internal longitudinal yes- 
sely of the teanchial sac. their erowding on the 
nartow tolds. the spherical vesicles embedded 
in the hranehial sac amd hady wall, the thick 
layer of circuhir musele, and the pnupillated pre- 
branchial area can be regarded as characteristec 
of this otherwise highly Variable single spuctes. 
The extent to which gonads are embedded in 
the body wall. and the exient to which the body 
wall is marked off inte areas probably indieutes 
more mature specimens. 


Palycarpa macniara Hartmeyer, 1906, has a 
similar endocarp enclosed by the gut loop. and 
ihe sume type of vesicles embedded in the 
hody wall, Tt es distinguished from the present 
species, however. hy the weaker musculature 


which also distinguishes 1 tram the West 
Indian species PF. ehiecia “Traustedt. 

PL pedunculata is The most common ascidian 
in St. Vineent Gulf and generally both greenish 
and reddish brown specimens occur. Large 
bluck specimens were also taken from Seal 
Rock. West t. from Hallett Cove, und from 
Tapley Shoal. There is no apporent cerrela- 
lion hetween the Lype of environment and the 
colour of the individuals wt ¢uch Station. A 
cuse of genetic polymorphism im Ascidiacea 
has been described for Boltenia ovifera CL) 
(Ploush 1969). This dominant in the ascistian 
population of the Gulf of Maine, has colours 
ranging From while to crimson red in a single 
boul. and variations in lest texture aod in 
muscle band colour and thickness can be 
related to these coluur variations. It has been 
suguested that the species demonstrates genetic 
segregation of the ability of individuals co 
jiccumulite pizments, The situation in Paly- 
corny pedunenulata may indicate a similar gene- 
tie segregation, 


Family PYURIDAR 
Pyura scoresbiensis 1.5). 

Type Location: Off Semaphore: 18 m, in 

sparse Posidonia, 27.1.69 (Holotype: Soush 

Australian Museum, registration number 

E876). Further Recards: Of Tapley Shoal, 

ifm. 22m 

FIGS. 57-59 

Deseription: Ronnded heads on stalks of vary- 
ing length, somcumes thick and no Jonger than 
the head, but sametimes long and. narrow (up 
to 20 em). supporting a head § em long and 
3 cm wide. The heal is more or less cge- 
shaped with its greatest diameter basally hefore 
narrowing abruptly te the stalk. The apertures 
une both sessile, either side of « more ar less 
pointed projection forming the anterior apex of 
the head. The atrial aperture is slightly more 
posterior than the branchial apertare 

The test is thin, hare and tough with a dense 
sindy encrustation on the outer surface of the 
head and the stalk, The body wall is thin and 
semidtransparent with moderately developed 
fine and diffuse musculature, with muscle bands 
most closely placed around the anterior part 
of the branchial sac and siphon. 

The branchial tenwacles hive a large Hanged 
axis. fairly short prinvury branches. stumps 
secondary branches and minute tertiary 
branches and are not very bushy. The siphons 
are lined with long needle-like spines, closely 
set, up to O<75 mm tong. There are no sm- 


ASCIDIANS OF SOUTH AUSTRALIA 1 


cules in either the test or the body wall. Vhe 
Horsal tubercle is a simple U-shaped opening 
with both horns lurned inwards. "The dorsal 
lamina has pointed languets but is very short 
owing to the close-set branchial and atrial 
siphons and contracted dorsum, The branchial 
sac is delicate with 6 high, overlapping folds 
on each side of the body with up to 20 internal 
longitudinal vessels On the olds and only 2 or 
3 between, There are 4 to 6 stigmata in cach 
imesh, 


There ts a simple and fairly narrew eu Joop 
enclosing the gonad on the left. The gunad 
on the right occupies a corresponding position. 
‘There are very arborescent liver lobules in the 
Tegion of the stamach, The gonad may consist 
fan undulating ovarian tohe with fringing 
testis follicles along hoth sides with the testis 
ducts extending along the mesial surfyce of the 
ovary. In some specimens the undulations of 
the covarian lube extend out into pinnate 
branches with tess follicles around their 
eXtremities. These pinnate branches may sub- 
sequently separate off into separate polycarp 
sacs on either side of a ventral duct. The anal 
border is divided into 3 large shallow Iohes, 


Hemerks: Specimens demonstrate the develop- 
ment of the polycurp sacs of the pyurid wonad 
from the continuous tubular styelid type of 
vonad. All stages of this development can be 
observed if the specimens available and it may 
be that the condition of the gonad indicates the 
age of the individuyl, The stalk’of this species 
alsa shows great variation in length and thick- 
ness. Despite these variations the species is 
characterised by the relatively smooth west, sand 
encrusted, bur without twhereles or furrows. 
und hy the constant position of the apertures. 
The pasition of the apertures, on the upper end 
nf the head, fairly close together, with the 
branchial anal atrial openings on opposile sides 
of the apex, is unusual in a stalked species of 
the Aseuliacea. where. more gererully, both 
apertures are on the dorsal side of the head 
with the branchial aperture directed uown- 
wards, and the atrial epefure uppermost and 
directed upwards. 

The relationships of this species are indicated 
by the stphonwl spines, which resemble those 
described for Pye albunyensis Michelsen & 
Hartmeyer, 1928, from Oyster Harbour. 
Albany, Western Australia. In which apertures 
wre also separated by » cushion of test in the 
middle of the upper surface and in which the 
dorsal surface of the hudy is very much con- 
tractedl and the dorsal! lamina comsequently very 


short. Pypra albanyensis has. however, charac 
teristic papillae on the convex bonier of the 
scubre-shaped stem and primary branches of 
the branchial tentacles. 


Pyara curvigona Tokioka, 1967, From the 
Palao Js. is a similar Closely related <pecies, 
sometimes stalked, with & similar arrangement 
of endocarps, gonads and gut, The anus, how- 
ever, has many lobes and the Jong (2.75 mm) 
siphonal spines extend outside the siphons onto 
the !ohes surrounding the apertures, as in 
Myra vittert (present in this collection). The 
needle-like siphonal spines found in the present 
Species are noc found in the various forms of 
the Pyara puchydermatina group of stalked 
species. In a specimen from Tupley Shoal 
{Station @} there are barnacles growing around 
the branchial aperture. 


Pyura vitlaty (Stimpson). Pérés, 1449: 195. 
Tokioka, 1952: 134; 19534: 273; 967: 
202. Millar, 1960: 126. Kot, 1964; (42: 
1946> 300; 1969: 133. For further syno- 
nymy wid literature to the species in the 
Aulanuc and West Indies see Van Name 
1945. 32). 

Cynthia vittata Simpson, 1852) 230. 

Pyura jacatrensis. Kou, 1952: 273; 1954- 127. 

Millar, 1960: 125. 
New Records: Tapley Shoal, off Trowbridge 
Light. Previous Records: W, Aust. (SW. 
Aust.)—Kott 1952, Tas.—Kott 1954, Qld, 

-Kott 1964, 1966, Pacific (Arafura Seca) 
~-Tokioka 1952: (Palao  Is.}-—-Tokioka 
1967; (Japanj)—Tokioka |953a; Van Name 
1945, Atlantic—WVan Name 1945; Pérés 
1949) Millar 1960, Sub-antaretic (Mac- 
quarie I,)—Kott 1954, 1969; (Kerguelen 1.) 
—Kott 1954; (Mation I.)—Millar 1966. 


The species has a wide circumpolar distribu- 
tion in the southern hemisphere and extends 
north through the Indo-Malayun region to 
Japan, It is also found in the Auantic and in 
the Cartibhean (see Van Name 1945). 


FIG. 40 
Description: Only a single individual is avan- 
able, 3 om long with a terminal branchial aper- 
ture and the vtrial opening half the distance 
alang the dorsal surface. Both apertures are 
almost sessile. “The external surface of the 
testis rough and has sand and foreign pirticles 
adhering. ‘The siphuns ure lined with tong 
needle-like spines, 0.1 mm to 0.2 mm long, 
overlapping. These extend onto the outer sur- 
face of the siphons, cover the lobes bordering 
the siphons and extend ome the outer layer 
wf test, The spines have a slight widescence 


ao VATRICLA KOLT 


which confers on this ouler siphonal urea a 
arcenish tinge. The siphon is lined with red 
siripes i the preserved specimen, 


The test is thin, leathery and firm, The 
dorsal tubercle is a rounded cushion filling the 
peritubercular aren with u simple U-shaped slit 
With both horns turned in. The branchial 
tentacles are nor bushy and have only primary 
briunehes und very shert secondary branches 
The internal siphons are fairly long. Longi- 
tudinal muscle bands radiate fram both siphons 
but do not extend very far down the bady on 
the Jeft. Circular muscles torn a Fairly irre- 
gulor network over the right side of the body. 
hecoming more sparse posteriorly. They are 
practically absent from the posterior hall of 
the hody on the lefe side, over the gut loop, 
The branchial suc is fairly delicate. It has 
18 tnternal Jongitudinal vessels on cach folk! 
and 4 between. There ate 6 stigmata per mesh 
Phe gut forms the usual logo enclosing the lett 
gonad, The anal border is smooth and bi- 
labiute, The gonads consist of the usual cen- 
trol ovarian tube with pinnyte branehes on both 
sides terminating in polyearp-like sacs,  Rndo- 
carp-like tissue 3s present on the free surface 
of the gonads where it is broken up into lobes. 


Remarks: Vhe synonymy of this widespread 
species has been very confusing owing ts) the 
variation in the length of the siphonal spines 
and the variation in the condition of the anal 
border, 11 appears. however, that Sluiter’s spe- 
cies from Indonesiit ind Northern Australia (P. 
jacutrensis). with very much smaller siphonal 
spines that do not extend onto the outer surface 
of the apertures, may he a distinct species Jes- 
pile the spines of intermediate length that are 
present in specimens from the Palau Istunds 
{Tokioks 1950; see Kott L9711. Pyura cvrvi- 
noma ‘Vokioka, 1967, trom Palao Is., 1s un 
wither closely related species in which the very 
long (2.75 mm) siphonul spines extend onto 
the outer surface of the apertures, In Pynra 
alhanyensis Michaelsen & Hartmever. 1928, 
and Pi xcoresbiensis nap. the siphonal spines 
extend up to 0.275 mm, only slightly Jonger 
than the present species. However, these 
siphonal spines do not extend onto che outer 
surface uf the apertures. 


Hyura icregularis (Herdman), Kott, 1952: 271 
Millar, 1963; 739: 1966; 370, 
Cwuthia ireecularis Herdman, 1881: 60; 1882+ 
141 
New Records: Tipara Reef, off Beach Hut, 
T ke off Port Vincent, upper St. Vinoent 
Gulf. off Grange, off West Beach, off Glen- 


elz, Port Nogtiunga, Aldinga “drop-oll". 
Cunchalings Head, Previous Records: S, 
Aust. (Quter Harhour), Vic, (Port Phillip 


Ray)—Millar 1963. 1966. Tas. (D’Entre- 
casteaux Channel)—Kott 1952. NWSW 
| Port Jackson)—Herdman 1852. The spe- 


cits fas nol prewously bean taken ih waters 
of less than 25 m in depth 


Deseviptior; Living specimens are red. orange 
tu light fawn. Externally the test is very hurd. 
leathery and wrinkled and thickened inte 
small oelugonul plates. There are also wart- 
like protuberances, especially anteriorly. 

Individuals are usually cluniped together in 
light aggregates und the shape of the body is 
consequently very icmegulan The maxim 
body length is about 2.5 em. Both apertures 
afe present af the end of EGuurly long siphons 
which are zenefally oriented away from one 
another. The test is very strong with internal 
longitudinal art outer circular muscle bands 
ws in all species of Pyuridue. 

Delicate cup-shaped scales, 0.02 mm long. 
line the siphons, There ure 15 branchial ten- 
tacles with short sparse primary branches wnd 
minute secondary branches. The primary 
opening from the neural gland is U-shaped 
with horns turned i er out. The dursal 
tubercle is blister-like and there is often an 
uccessory openiny [rom the neural gland, The 
tubercle is not always longitudinally attenua- 
ted. however the peritubercular are 15 always a 
very deep V-shape und generally the tubercle 
does extend dawn into it. The neural ganglion 
is especially long, extending most of the dis- 
tance atone the dorsal lamina, The dorsal 
Juminu has a dowble row of Languets, These 
are fine anil pointed. closely set on the left. 
und on the right they are stouter and more 
sparsely urrunged. 

‘There are from & to 10 branchial folds un 
each side of the body with ahout 12 longi- 
tudinal vessels on the folds and 2 between. 
There are 6 to 8 stigmata in euch mesh crossed 
by parastigmatic vessels, The gut loop is 
simple and curved and encloses the left gonad 
which is subdivided into 15 1 20 separate 
polvearp sacs arranged on either side of central 
male and female ducts, There is a corres- 
ponding gonad on the right. 


Remarks: This species resembles very closely 
the Antarctic species Pyura discoveryi Herd- 
man (see Kott 1969). ‘Che tough, wrinkled 
external test with embedded polygonal thicken- 
ings is alsa reminiscent of the Antarctic Pyare 
squsniata Herdman although the polygonal 


ASCIDIANS OF SOUTH AUSTRALIA 3y 


seules und the body shape of P. squameta are 
more highly specialised than in either P. ass. 
covers? or in the present species The bran- 
chijil tentacles with their sparse branches und 
the long siphons ure alsa similar to those of 
P. discoveryi and it is possible that the protec- 
lion afforded the individual by these Jone 
siphons may be assncigied with the absence of 
the more bushy tentacles usually found m this 
uchus. 

The individuals are never very farve and 
their leathery test and hahit of occurring in 
ageregates suggests a species udapled Jor yery 
turbulent conditions. Che present records do 
not support this, however as they are vither 
fram Offshore Benthic locations in St. Vincent 
Gulf. ar from reefs in sheltered coastal loca- 
ions. 


Pyora australis (Quoy & Gaimard) 5.sp. ausira- 
fis Quoy & Gaimard. 
Aseidia vastralix Quoy & Guimard, 1234: 614, 
Priva australis €. rypieq, Kou, 1952: 268 and 
synonymy, 
Pivre australis, Millar, 963° 739. 
New Records: Vipata Reef, Tapley. Shoal. 
near Murion Light, off West Beach, off 
Rroadway. off Hallelt Cove, off Yankalilla 
Baw West LL, N.W. of Robe. Previous 
Records: W. Aust. (Geraldton to Albany) 
—Quoy & Gaimard 1834; Michaelsen & 
Hartmeyer 1928: Kou 1952; Millar 1963. 
Vic, (Weslermport, Flioders}—Quuy & 
Gatmard $8342 Koi 1952. Tus. (D’Entre- 
casteaux Channel, Tinderbox)—Kott 1952. 


FIG. 61 


PDescripiion: Specimens of all sizes up tu a 
maximum of 4 cm long head with a stalk of 
30 cm. ‘The lest is usually without foreign 
bodies adhering, though in one specimen there 
are some cirtipedes growing on the stalk. The 
surface of the test is marked with variable 
Jongitudinal furrows pnd ridges but is some- 
times almost smooth. In preservative the 
specimens are pinkish-fawn, although living 
specimens. ure usually dark red and, Occa- 
sidually, yellow. Both apertures are close 
together on the dorsal surface, the atrial uper- 
ture directed upwards and the branchial aper- 
ture directed bisully.. The Jobes of the atrial 
uperture wre clearly continuous with the ridges 
in the dorsal part of the test 


There are stellate spicules of about 0,02 mm 
diameter with 6 rays i optical transverse sec- 
tion in the body wall, and the siphons are lined 
by conical spines of U.02 mm maximum height 
from buse to apex, 


The bronchial sac, gut loop and gonad ure 
as previously described and there arc 18 long 
fisttened characteristic lobes fringing the anul 
border (see Kott 1952). 


Remarks: Nothing can be added to previous 
descriptions of this constant species which 
appears. to occupy a wide Tange of conddirions 
in exposed to sheltercd locations from Geralu- 
ton, in Western Australia, to Flinders in Vic- 
loria. It is common in wave besten areas 
from the low water mark to 22 nv. 


Pyura spinifera (Quoy & Gajimard). Kot, 1952, 
269 and synonymy. 

Aridio spintfera Quoy & Gaimard, 1834; 6) 7_ 
Cyethie multiradicata Herdman, 18¥9: 30. 
Now Records: Upper St, Vincent Guif, off 
Hallet. Cove, Aldinga. Previous Records: 
Wo Aust. (Albany)—Quoy & Gaimard 
1834. Vie. (Buss Strait)—Michi#elsen 1905; 
Heller 1878. NS.W, {Port Jnckson. Port 
Hacking)—Werdman 1891. 1899; Kott 

1952. 
FIGS. 62, 63 


Description. Specimens with head to 8 cm tong 
am 4.5 em wide. Stalk is of very variable 
length. maximum 20cm. Externally the test 
is smooth without longitudinal burrows. but 
with characteristic tubercles, varying in their 
density, and sometimes. especially in Jarger 
specimens, absent ultogether, The head is often 
completely enveloped by an investing sponge 
which in specimens from olf Hallett Cove hus 
been noted in the field as yellow. 


Minute overlapping scales, 0.05 mm tmani- 
mum length from posterior patt of the base 
to their apex. line localised areas where thick- 
ened lobes of the test project into the siphons. 
Otherwise. there ure no spicules in the test or 
in the hody wall, There sre 7 branchinl folds 
on cither side of the body wall in the larger 
specimen but only 6 on cach side in averuge- 
sized to smaller specimens. There are abou 
25 branchial tentacles alternating with rudi- 
mentary tentacles. The larger tentacles have 
regular pinnate primary branches with seoon- 
dary branches and minute tertiary branches 
and are very bushy. The dorsal tubercle has a 
double coiled opening. both horns coiled 
inwards anc the inner spirals of cach coil are 
slightly convoluted. There is <i short dorsal 
lantina with pojnted Iqnguets, 

There are yp to 30 internal longitudinal ves- 


sels on the folds and 2 to 3 between, In larger 
specimens the under sides of all the mujor 


40 PATRICIA KOTT 


65 


4a ah 
hy 


8 ny 


y —4 


Figs. 57-59. Pyura scoreshiensis. Fig. 57—Individual (off Semaphore, 18 m). Fig. 58, Siphonal 
spines. Fig. 59—Gut and gonads. 

Fig. 60. Pyura vittata. (Tapley Shoal, off Troubridge Light, 17 m). Siphonal spines. 

Fig. 61. Pyura australis. (Haljett Cove, 8 m). Spicules embedded in siphonal lining, and siphonal 
spine. 

Figs. 62, 63. Pyura spinifera. (Upper St. Vincent Gulf, 10-11 m). Fig. 62.—Dorsal tubercles. Fig. 
63.—Papillac from inner body wall. 


Fig, 64. Microcosmus nichollsi. (Off Yankalilla Bay, 20m). Siphonal spines and scales. 
Fig. 65. Micracosmus squamiger. (Off Semaphore, 18 m). Siphonal scales. 
Fig. 66. Microcosmus stolonifera. (Port Noarlunga, 5-6 m). Siphonal spines. 


Figs. 67. 68. Crenicella antipoda. (Yankalilla Bay, 12-20 m). Fig. 67—Dorsal tubercle. Fig. 68.— 
Inner body wall showing gonads ani gut loop and heart on left and right respectively. 


en TEENIE as 


blood vessels and the transverse vessels (but pointed languets to form a fur-like covering. 
not the parastigmatic vessels) support minute These projections also cover the gonads and 


ASCIDIANS OF SOUTH AVSTRAL TIA él 


the whole innec surface of the body wail 
extending into the buse of the atrial siphon 
although here they are reduced in density. 


The gut forms a narrow curved loop enctos- 
ing the left gonad. The right gonad forms a 
corresponding curve on the right side of the 
body. “Vhe anus is bordered by 12 shallow 
lobes. There is a mass of orange arborescent 
liver lobes. In larger specimens there is a 
blister-like structure on either side of the atrial 
opening. extending into the curve of the gut 
loop and into the curve of the gonad on the 
left und tight sides of the body respectively. 
‘This also has a fur-like surface formed by 
dense. small, pointed projections. The inner 
cavity of this blister-lke organ is continuous 
into the lumen of the virial siphon and, pre 
aumahly, iF swollen or distended could occhide 
the lumen of the siphon. ‘Phere are also two 
flaps of tissue, anterior abd postenor to the 
atrial Opening to form an wtrial velum. 


Remarks; This distinctive species, in which 
varialion in exterial appearance involves only 
the nuniber ef tubercles on the test and the 
length of the stalk has, in Si, Vincent Gulf, 
only been taken from fairly sheltered situa- 
tions, Other records, however, suggest that 
ithe species could occupy greater depths in off- 
shore situations from which it was uprooted 
omly with turbulence occurring during storms. 
The large head supported on the thin but tough 
stalk does not appear te favour yery rough 
conditions, although it could he an advantage 
in locations Where there is steady current flow 
or surge. 


Halocynthia hispida (Herdman). Kott, 1968: 77 

and synonymy. 

Cyathia hispida Herdman 1882: 146, 

Holeeyuthia cactus. Vasseur, 1967; 144. 
Mew Records: Tipara Reef, Tapley Shoal, 
hear Marion light, off Beach Hut, Port 
Vincent. upper St. Vincent Gulf, off Outer 
Harbour, off Wesr Beach, off Glenelg, off 
Pert Stanvac (“The Barges"), Aldinga, 
Carickslinga Heat, off Yankalilla Bay, 
Rapid Hed, Previows Recordy: See Katt, 
1968, 


Remarks; This species upparently occupies a 
wide variety of conditions but generally favours 
sheltered bays or estuaries (see Kott 1968. for 
description and further discussion of this and 
related species). 


BReriuania momus (Sayigny) Michaelsen, 1919: 
30 and synonymy, 


Cynthia mimus Savigmny, (HIG: 143, 

Pynra mons F Rvarhirensiy Michaelsen, 
1949" 44, 

? Pyura moms F petunet Michelsen, 1419: 


Pynra munis f. 

1949: Sa. 

Pynrad mmomhes f. galei Michnelsen & Hurt- 

meyer, 1927: 194, 1928; 443, 

Pyura momus Savigny t. grantis. Michaelsen 

& Iarimeyer, 1928: 44. 

Heedniania momus. Van Nanie, (9452 344 

Herdmiania niamis (. culei. Koll, 1952. 381, 

Tokioka, (961; 132; 1967: 205 

Terdrnania momus £, grandis. Katt, 1952: 

279, 1964: 142; 196A: 301. Millar, 1960: 124: 

1963; 740; 1964: 374. Tokioka, 1949; BI: 

1952: 137: 1953a° 277; 19A7: 2G. 

 flerdmania momus f, crrvere Kou, (952: 

282; 1964: 143. 
New Records: (“grandis” type)— Tipara 
Reef, off West Beach, off Glenelg, Caricka- 
Jinga Head, N.W. of Robe. (‘gale type) — 
Goose |. upper St. Vincent Gulf, Aldinga 
Reef, West b, Wright |. Previous Records 
(“grandis typey: W. Aust, (Fremantle to 
Albany )—Michaelsen & Hartmeyer 1928: 
Millur 1963. Vic. (Port Phillip Bay, Wes- 
ternport)— Millar 1960, 1963, ]966. N.SAV 
(Port Jackson)—Heller 1878; Herdman 
1882; von Drasche 1884; Vokioka 1967: Mil- 
lar 1963, Old, (Bowen)]—Kott 1952 Indo- 
nesia (Off West Irian—Herdman | 886: Ara- 
fura Sea)—Tokioka 1952. Japan- Vokioka 
1949. Pucific ¢Piji Is.)—Herdman 1882; 
Palao, Tahiti) —Heller 1878. Indian Ocean 
(West Indiin Ocean)J—Michuelsen 1908: 
Heller 1878: (Red Seaxj)—Michuelsen 1919: 
Savigay 1816; (Dar<s-Salaam)—Michael- 
sen 1905; (Ceylon) Herdman 1906. Africa 
(Cape of Good Hope, Simons Bay}—Herd- 
man 1832. West Indies (Jamuaica)— 
Heller 1878 (“gale type) —W-. Aust. 
{Shark Bay, Point Charles, Dirk Hartog 1) 
—Michaelsen & Wartmeyer 1927, (928. 
Tas. N.S,W. (Port Stephens), Old. (Bowen, 
Nelson’s Bay)—Kott 1952, Pacific (Mela- 
nesia)—Tokioka 1961; {Marianas Ts.)— 
Tokioka 1967; (Japan)—Tokiokn 1967. 
(For records of specimens recorded as 
“pallida” form, see Van Name 1945). 


Michaelsen (1919) has considered, in some 
deturl, the distribution of all the forms of this 
species. Apart from certain forms represented 
by single records, many of the ranges Overlap 
and no separate geographic runges can be 
assigned. The range of the species, repre- 
semted by the range of the form pallida, for 
which there arc most records, is civcum- 
tropical, and extending south tu the Cape of 


vevnplandta. Miclaélsen, 


42 PATRICIA KROTT 


Good Hope. Forms from the south cuaast of 
Australia have been described as forma 
erandis, ‘This form is not, however, cistinct 
from f, pallida (see below) and it ts doubtful 
whether there is justification for separating any 
of the specimens assigned to the species. Their 
morphological variations are most probibly 


indicative of different stages of maturity, 


Description: Vhe distribution of the severul 
forms. L, manus t. grandiv. H. momus F. 
pallida and MH. morrtuy f. galei, overlaps and in 
the present collection all forms have been taken 
from the sume locution and it is apparent that 
H, momas £, grandis with an opaque 
whitish test. a convoluted dorsal tubercular 
opening and with testis follicles covering the 
ovary. represents mature individuals of a 
species in which the juvenile specimens have a 
jranspurent to translucent Les! with the testes 
follicles arranged révularly around the peri- 
phery of the ovary (f. gale’), Sometimes in 
intermediate sized specimens the eyarian Lube 
undulates along its length und the testis fol- 
licles may remain close lo the ovary (as des- 
arihed for f pallida; Van Name, 1945), In 
other specimens in this collection (3 km off 
Glenele) the testis follicles form an even 
border around an urea in which the ovarian 
tube ix undulating, In the smaslest specimens 
the anal lobes are rudimentary, later develop 
inlo even fingerl{ke Nattened Jobes, which 
become less regular and may be absent in 
larger specimens. but are sometimes present in 
two clumps at either side of the opening. 


Remurky: {tis apparent from the present col- 
lection thal the pele, grandix and pullicde 
forms of this species represent different stages 
of maturity of a single species. The relution- 
ship of the present forms, in which the ovaries 
undulate with the testes follicles which some- 
times cover it. to Il. momus f. typica Savigny 
(CH, momuy {. chrrvat Kott, 1952; 1964) in 
which the testes follicles are arranged in an 
undulating line along the ovary, is problema- 
tical. However, it is probable that the undula- 
tion of the ovarian tube could have forced the 
testes follicles int at similarly undulating line. 


Microcosmus nichollst Katt, 1952; 290, 


New Records: OF Beach Hut, 1 km off Port 
Vineent, off West Beach. off Hallett Cove, 
Aldinea, Carickalinga Head, West 1, Wright 
L Previons Records: Vic, (Flinders) —Kart 
y4s2 


FIG, 64 


Deseriptian: Vest generulty thick, whitish and 
coriaccous with pinkish colour around siphon 
but sometimes tough and almost leathery 
externally with rounded ridges or thin, stiff. 
rough and embedded with sand. uneven and 
marked by horny scule-like areas. Externally 
both apertures are sessile and close together on 
the upper surface. each surrounded by raised. 
rounded projections of the fest. Posteriorly 
the test may be produced into root-like pro- 
cesses. There ts a network of longitudinal anu 
rectanuular muscles. 


The siphons ute long and the siphonal mus- 
culature is especially strong, Outer circular 
sphincter muscles surround the base of each 
siphon and the longitudinal muscles extend 
ucross the body bur ary absent. from the region 
over the gut Pojnted conical spines and 
smaller spines and more numerous scales line 
the siphons. There are sometimes calcareous 
spicules embedded in the body wall and in the 
Jentacles and branchial sac, Branchial ten- 
tucles have primary. secondary and minute ter- 
tiary branches. The dorsal tubercle is 
U-shaped with horns Iurned in. The dorsal 
ganglion is elongate, half the length of the 
wide. plainsedged dorsal lamina. There is a 
pronounced branchial vellum, On euch side of 
the body wall there are high overlapping 
branchial folds with uj to 20 jnternal longi- 
tudinal vessels on the folds and 1 to 3 between, 
There are about LO stigmata per mesh. hetween 
the folds, crossed hy purastigmatic vessels, The 
wut forms u simple closed and nurrow loop 
around the ventral border of the body enclos- 
ing the terminal lobe of the gonad in its loop 
The descending limb is crossed by gonad. 


‘There is a stomach enlurgement obscured by 


liver lamellae which arc smaller at the pyloric 
end of the stumach. Minute finger-like pro- 
jections Eram the surface of the liver lanictlae 
give it a furry appearance. The anus is bor- 
dered by 12 rounded lobes. 


On the right, the gonad curves around the 
ventral border und on the left curves inte the 
loop of the gut just distal te the Tiver lobes 
The vonads are broken into 2 rounded clumps 
on the right and 3 on the left, often covered 
hy enducarp. 


Remarks: “The small siphonal scales and the 
gonad across the gut loop, together with the 
whitish and more gelatinous test of the 
smaller specimens, distinguish the species fram 
M. stolonifera, 


ASCIDIANS OF SOUTH AUSTRALTA 43 


Microcosmus squamiger Michaelsen, 
Microcesmmus claudicans subs sp. syuamiger 
Michaelsen & Hartneyer, 1928: 405, 
Microcosmus exasperaius sub. sp. australis, 
Michuelsen, 1908; 272; 1978! 69 (ine part, 
excluding MW, aitstraliy Hérdman, and MM, 
rantgey’ Herdmatr), 

New Revoreds) Tipara Reef, off Semaphore, 
off West Beach, off Glenelg. Previous 
Revords: W. Aust. (Shark Bay to Albany) 
—Michiaelsen & Hurtmeyer 1928, N.S.W. 
(Sydney)——Michuetsen 1908. Old. (Bowen, 


Rockhampion)—Michuelsen 1908, Red Sea 
—Michnelsen J9]8. 
FIG, 65 


Deseriplior; Small. Iwathery, pinkish speci- 
Mens, aggregated together, The surface of the 
lest is raised ino ridges und mounds, The body 
wall is very muscular. The dorsal tuhercle is 
large with a double spiral opening. There are 
the usual 8 branchitl folds on each side of the 
body and the left gonad crasses into the gut 
loop. The gonad ov each side of the body is 
divided into 3 clumps. There are close-ser 
liver liimellae, Closely set curved scales 0.112 
mm Jong line the branchiil siphon, 

Remarks: There has been some contusion 
between Af. exasperatus, M, sustralis, and the 
Present species, all common around the Aus- 
tralian coast and all demonstrating a fairly 
wide diversily in external appearance. The 
reddish colour and agercgated habit, the large 
number of tough branchial folds, the deeply 
curved gut loop and the gonad crossing into 
the gut loop, are chiracters shared by all three 
species, Microcosmius squartiger is dislin- 
guished by flattened scale-like siphonal scales, 
while both Microceymus australis Herdman 
and Mf. exesperatus have pointed spines. 


Microcosmus stolonifera Kutt, 1952: 291, 
New Records: Tipara Reef. Port Noarlunga. 
Previous Record: Tas. (Tiny 16, east coust) 
—Kot 1952. 

FIG, 66 

Description, Only two specimens are available. 

They ure very irregular externally, and pos- 

teriorly are produced into rool-like processes. 

The upertures are on siphons of yarinhle 

length. turned away from one another and, in 

the largest specimen available (2 cm greatest 
dimension! the siphons are especially long. 

The lest is very tough, hard and leathery, There 

are large (about 0.1 mm) pointed spines, 

arfanged in fairly regular horizontal rows. lin- 
ing the siphons. The branchial tentacles are 
bushy. The branchial sac has 7 high and deli- 


cate overlapping folds, with a single internal 
longitudinal vessel in the interspace, The suit 
Forms u narrow curved loop with the usual 
elongate liver lamellae with short finger-like 
papillae from its surface. The gonuds form 
a single rounded mass in the curve of the gut 
Joop on the left but do not extend inte the 
primary gut loop. On the right there may be 
& corresponding single rounded anasx or thir 
Tight gonad is somelimes divided into two 
rounded lobes joined by the central ducts. 
Remarks: The vest of this species Ts harder and 
Joys reyular than all other species of this venus, 
fl is further distinguished by the long siphonal 
spines, the large rounded gonad that does not 
develop inside the gut loop, and the high deli- 
cate overlapping Folds of the branchial sac. 

je does not appear to be a very gommon 
species and the only two recoms ufe from 
the southern coast of Australia. However. the 
tough and roughened test, forming a very 
strong attachment. causes the species ii he 
inconspicuous und difficult to collect. 


Mierocosmus helleri Herdman, 1880: 54; 1882: 
131. Slutter, 1895; 184, Hartmeyer, 1919: 
19. Michaelsen & Hartmeyer, }928; 397, 
Kott. 1952: 292; 1972; 12. Millar, 1463: 
742, 

Mictoeasnins eoanus Michuelsen, 19182 12, 
New Records: Tapley Shoal, olf Beach Hut 
(J km off Port Vincent). Previews Records: 
W. Aust, (Cape Jaubevt to Fremantle )— 
Harumcyer 1919; Michaclsen & Harimeyer 
1928, Koti 1952; Millar 1963. Old, (Great 
Barrier Recf)—Kou 1952: (Torres Strain 
—Herdman )882. Malaysia—Sliuiter 1895 
Portugese East Africa (Delagoa Bay)— 
Michaelsen 1918. 


Description: The single spherichl specimen 
from Tapley Shoal is 6 cm in diameter. This 
lirge diameter is contributed to by a } em thiek 
coating of sand held together by terminally 
branching and coalescing projections from the 
test to folm a thick dense layer enclosing a 
space around the body. This coating is inter- 
Tupted to form uw single opening above the 
apertures, ‘The specimen from of! Beach Hut 
is more typically rough externally and is a 
purple colour, The apertures are sessile, one- 
third of the body circumference apart. At the 
base of the branchial siphon there wre 3 flap- 
like projections. 

The body musculature is of the usual pyucid 
type with muscle bunds from each of the 
siphons crossing ane another on both sides of 


4 PATRICIA KOTT 


the body. There are very strong circular 
mniscles. circling each siphon, 

Branchial tentacles bave pnmary and secon- 
dary branches. and wide, flat, oveinbraneous 
extensions from their anterior or concave 
border, The dorsal lamina is plain. There ure 
6 high, overlapping folds on cach side of the 
body with up to 18 internal longitudinal ves- 
sels on the folds and 3 between, The gut 
forms the usual long, narrow ultenuated loop, 
typical of the species, and the proximal lobe 
al the 3 lobed left gonad is accommodated in 
the apen pole of the otherwise closed gut loop. 


Remarkys The tough flap-like projections in 
the branchial siphon sometimes appear as 
cones, These structures, together with the gut 
loop und branchial sac, distinguish the species. 


The sandy coating has not been described 
previously for this species, but has been des- 
eribed For Pyura concellate Brewin from New 
Zealind (see Kott 1971) and for Pynre tunica 
Kott, 1969 from the Anturetic. ‘This condi- 
tion demonstrates the versatility of the ascidian 
test which in this specimen responds to the sub- 
strate by growing out to entangle sand grains 
as there is no firm substrate onto which it can 
directly be fixed. 

Ctenicella antipoda n-sp. 

Type Loevatty: Of Yankalilla Bay. at 12 to 

20 m (2 specimens): in Amphibolis com- 

munity With limestone outcropping occa- 

stonally. Holotype: South Australian 

Museum (reg. no. E877). Further Recorst: 

Tipurca Reef. 

FIGS. 67, 68 


Heseription. Specimens are up to 10 cm long, 
slightly dorso-ventrally fluttened. Externally 
they are very irregular and covered with 
nodules which ali protect the sessile apertures 
on the dorsal or wpper surface. The test ix 
up tw 1.5 em thick. gelatinous but entirely 
Jmpregnated with sand so that it is bard and 
rigid. {1 is sometimes produced into a ridge sure 
rounding the siphons, There ure hard brown 
pupillue around the sessile apertures but there 
are no spines lining the siphons. 

The body musculature is strong on the upper 
half of the body with longitudinal bunds radia- 
ting from the siphons and inner circular bands 
around the siphons and at their base. How- 
ever, on the lower half of the body the muscu- 
lature is almost entirely absent and is repre- 
sented by two vertical rows of very short 
parallel bands, 


There are 15 lurge compound branchial ten- 
tucles with primary, secondary, and minute 
tertiary branches wlternuting wilh rudimentary 
tentacles, “he dorsal tubercle is at the base of 
the tentacles anterior to the Voof the per 
tubercular arca. The opening is a double 
spiral slit turned to the left. The dorsal famina 
is very short and has close-set slender. pointed 
lunguets, 


The branchial sac has 6 high, overlapping 
folds on euch side of the body. widely spread 
at their base, Longitudioal vessels are urranged 
as follows: 

DL Be MISSI S(AS AAAI FLI4) 2153 EL 


There ate about [2 stigmata in each mesh. 
They are rectangular. and crossed by para- 
stiginatic Vessels. ‘The meshes are wider than 
Tong and there ts no sign of itregulanty in the 
stigmatit which do not coil nor form infundi- 
bula, 


‘The gut forms a ourrow. closed and decply 
curved loop with branched liver lobules extend- 
ing along the inside of the gut loop for its 
whole length. ‘he liver is spongy with shan 
rounded finger-like papillae projecting: from its 
surface. and supporting cissie between the liver 
lobules, 


The intestine is filed with mud. ‘The anal 
Sorder has about 30 or more rounded lobes. 
On the tight side of the body there is u long 
curved hypertrophical heart in the posilion 
occupied by the kidney in Molguliduc. There 
is a single gonad on cach side of the bacly 
piurallel to and lying against the long conspicu- 
ous heart on the right, and on the lett extend- 
ing parallel to the descending line of the pri- 
mary gut loop, ‘The lef gonad descends into 
the secondary gut loop where its short ducts 
turn dorsally toward the atrial aperture. The 
ovary is central and tubular. while the espe- 
cially small pyriferm tess lobes extend into 
folds in its wall, giving the appearance of being 
embexitled in the ovary. Th one of the speci- 
mens from Tipara reef the gonads. ure mir 
malure and groups of very minute testis lobes 
are arranged around the upper ind outer sur- 
Pace of both sides of the ovary. Vasa efferen- 
tin from each group of follicles join together 
to open into the vas deferens along the 
medtun surface of the ovary. 


Remarks: Ctenicella Lacaze Duthiers (Type 
Species: Ctenicella appendiculata (Heller). 
From the Mediterranean), has few known 
species, although a number of Malgila spp. 


ASCIDIANS OF SOUTH AUSTRALIA as 


have been crroncously ascribed to it, The 
genus is characterised by the presence of dorsal 
languets, straight stigmata, a kidney on the 
right. and the left gonad outside the primary 
gut Joop, Inaddition to the type species which 
is. distinguished by its long recurved siphons, 
Crenicella undulata Tokioka, 1949, from 
Japan, has a posterior stalk and a folded 
stomach, 

Hurtmeyeria Ritter was also thought to be 
intermediate between Pyuridae and Molgulidac, 
with pyurid branebial sac, siphonal spines, a 
smooth dorsal lamina and the left gonad partly 
in the gut loop (as in certain species of Micra- 
casmes). Monniot (1969) has shawn, how- 
ever, that what was thought to be a kidney, is 
in fact un hypertrophied heart and that Hart- 
reverie is without doubt a pyurid genus related 
lo Micracasnaty and with a liver similar to 
that of Hulocynthia with longitudinal plications 
proximally and branched tubules distally, Aart- 
meyerla differs trom the present species in 
us smooth dorsal lumina and siphonal spines, 
and in the position of its left gonad which 
crosses into the gut loop. It is probuble that 


the kidney, which has been deseribed for 
Crenicella undulata and -C'. appendteulata is, in 
fact, an hypertrophical heart, as described for 
Harimeyeria and us demonstrated for the pre- 
sent species, 

The identity of Crenivella undilata Vokioka 
is puzzling as it has dorsal Janguets and the 
gonads on the left and right respectively in the 
usual position for the genus, outside the gut 
loop and adjacent to what has been described 
ig an excretory argan, However, the stomach 
uppears tO have proximal glandular folds and 
distal arhovescent lobes as described for Hary- 
meveria and Halocyntiia and it has a Aari- 
meyeria type of stalk. Therefore, both Creni- 
cella and Hartmeyesia appear to be genera of 
the Pyuridae, distinguished from Pyura, Hula- 
cynthia and Micrecosmus by un hypertrophied 
heart. They appear to be distinguished from 
onc unother only by the absence of siphonal 
spines, the presence of dorsal Janguety and by 
the position of the gonad outside the primary 
gut Joop in Crenicelle spp, The télationships 
of these pyurid genera are shown in the follow- 
ing Table. 


TABLE 1} 

Camparinin of Characteristics. of the Genera of the Femify Puiridac. 

Pyura Halecynihia Ctenicella Horimeveria Micrecasmus 
Siphonal spines present present none Ppreseni present 
Derséllaniina — languets languets languets smooth amoath 
Liver tissue urburescent long foalisand — arboresceni leng folds and — stborescent 

lobes athorescent lobes : arhorescent lobes 

lobes (I species with lobes 


Gonads in primary cross gut loop 


But laop 


long folds and 
arbarescent 
lobes} 
autside gutloop cross gut laap 


trass gut loup 


ees 


Family MOLGULIDAR 
Molgula mollis Herdman, 1899: 54. Kott, 
1952; 298; 1964: 144. 
Molenly sydnevensis Herdman, 1899; 55, 
pian janis Kott, 1952: 295. Millar, 1966; 
4. 


New Record: Carickulinga Head. Previous 
Rerords; N.S.W.. (Port Jackson, Sydney )— 
Herdnaan 1899; (Twofold Buy) Kort 1952. 
Old. (Gladstone to Moreton Bay)--Kott 
1964, 
Description: Small, rounded, laterally Nattened 
specimen of 0.6 cm diameter. The apertures 
are present anteriorly in a depressed, sand- 
free area of test, surrounded by sandy pro- 
tuberances and hairs fram the thin test. 


The dorsal tubercle is oval with a longitu- 
dinal, more or less S-shaped slit, The neural 
gland is conspicuous beneath the tubercle, 


The branchial sac has 7 folds on each side 
of the body with only 2 internal tongijudinal 
vessels along the top of each fold. Stigmata 
coil to form infundibula projecting into the 
fokls and subdividing into two in the summit 
of the fold, Between the folds there are some 
interstitial stiymatal coils but no primary 
infundibula, The spirals of the primary coils 


ate interrupted in their median longitudinal 


and transverse planes and their arrangement. 
especially at the base of the spiral between 
te folds, is obscured, 


an PATRICIA KOTT 


The gonads are flask-shaped and the [(estis 
follicles form. circle around the proximal end 
of the ovary, with iw connective from the centre 
pf this circle us previously described (Millar 
1966). 


Remarks: The species is characterised by the 


small number of longitudinal vessels on one 
side of the branchial folds. There is some 
variation tn the development of the hollow 


extensions of the test which Kott (1952) had 


thought distinguished M. janis, Tt is clear 
however, that the species is synonymous wilh 
MM. mallty. 


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Appendix I—Siation List 
A. ROLGH COAST SUBFORMATION 


West [scann; on eranite usually on vertical Fiees 
OT IM caves, 


1. Region A: rough [Shepherd & Womersley 
1970), depth indigaled for each speciey. 
Podoclavella eylindriea 25 na 
Leptaclinides rufus iim 
Suirylloides magnicnesuy 22-25 m 
Retryloides leachi bf 25 m 
Hoprylaides nigrum (2-20 m 
Oculinaria ausivalis, 12-25 m 
Cnemidocarpa etheridgit 25 m 
Polycarpa peduncilatih 16-25 m 
Pyura australis 1220m 
Micracosmay nickallsi Ze 
flerdmania mantis 16-22 tn 


Region B: moderately rough (Shepherd & 
Womersley 1970); depth IS m. 

Syeozow cerebriformis 

Palycitor gigaateum. 

Boirvilaidex leach 


re 


3, Region D: sheltered (Shepherd & Wormersley 
L970); depth 2-5 m; 27.044, 
Podocluvella cyligdrica 
Cystedves dellechtajed 
Synoicinm papillijerum 
Ofdernnam candiedum 
Didemnum moseleyi 
TridldemaAtm sxpiculatim 
Didemnum sp, 

Leproclinides rufus 
Borrylloides nigrum 


Wiunitty Istanp! roogh coast, strong surge: on ver- 
tical granite faces; depth 10 my 28.x7,66. 
Padoclavella cylindric 


5) PATRICIA KOTT 


Sy¥cozea cerebriformis 
Atapozog fantasiaia 
Polycitor afgantenm 
Eudixtenis renieti 
Didennunt candida 
Lentovlinides rufis 
Phallusia depressitseulit 
Ascidia sydneyensis 
Botrylloides leachi 
Borryviloides nigrum 
Oculinaria australis 
Cremitocarpea etheridgil 
Polycarpa pednncalata 
Microcosminus nichollsi 
Herdmania mors 


KinG Beaci, Encounter Bay: under boulder on 
intertidal reef. 
Corella enmyota 


Nora Creina Bay, near Robe: on roof of cave, 
strong surge: depth 1) my; 11.41.67. 

Eudistoma sp. 

Pseudodistoma cercurm 


24 KM NorTH-west oF Rose, Svuth Australia: on 
aeolianile; slight surge; attached to red algae; 
depth 40 m.: 20-x7.68. 

Pyara australis 

Heérdmania moms 


B. SHELTERED COAST SUBFORMATION 


Orr HALLetr Cove, on reef: rocky boltom: depth 

8 m: 26.x11,66. 
Podoclavella cylindrica 
Distaplia virtdiy 
Sycozoa cerehriformis 
Polycitor giganteum 
Aplidinm pliciferum 
Leptoclinides rifts 
Echinoclinum verrilli 
Rhadosemea tarcicum 
Corella cumyota 
Phaflusit depressinseula 
Ascidia thompsani 
Ascidia svdnevensiy 
Polycarpa pedunculaia 
Microcasmus nichollst 


Insipc Port NoarRitinGA BREF: moderate surge, 
in caves or on vertical faces; depth 2-5 mm; 
20,x1.66. 

Podoclavella evlindrica 

Distaplia viridiy 

Ritterélla herdmania 

Synoicinm papilliferunt 

Leptaclinides rufus (sometimes investing Pyura 

irregilaris and Microcosmus stolontfera) 

Ascidig sydneyensis 

Botrylloides feachi 

Stolonica australis 

Polycarpa pedunculata 

Pyura trregilaris 

Microcosmus stolonifera 


ALDINGA, Reer at “Drop-orr’: racky bottom; 
slight surge: depth 10-25 m; 12.xii66. 

Podoclavella cylindrica 

Palycitur giganieum 

Didemnum lambitum 

Didemnum patulam 

Palycarpe papillata 

Polvcarpa pedunculata 

Pyura irrepularis 


Pytira spinifera 
Haloexnthia hixpice 
Herdmenia monius 
Microcosmus nichollst 


CARICKALINGA HoaD: in caves and on vertical rock 
faces, moderate surge: depth 5-h m; 18.11.67. 

Clavelina haudinensis 

Disteplic virtelix 

Sycozea cerchrifarmisx 

Didertnum moseleyi 

Ascidia thompson 

Botrvilotdes nigrum 

Palycarpa podunculiter 

Pynra irregularis 

Herdnumia morins 

Hadlocynihia hispida 

Microcosmay nichollsi 

Moleula mollix 


Rapip Heab: on vertical faces dnd under ledges: 
Slight to maderate surge; depth 10 m; 25.iv,66, 

Clavelina baudinensis 

Polysyneraton erhiculum 

Léptoclinides rufus 

Botrylloides nigtum 

Polycarpa pedunciulata 

Halocynthia hispida 


OFFSHORE. BENTHIC LOCATIONS 


Goose [,, Spencer Gulf: on rocky bottom, depth 
3-S m:. 1-x.66. 

Didemnum moselevi 

Herdmania mamus 


Treara REEF. Spencer Gulf: 
1. on travertine vertical faces and under ledges: 
depth 6 m: 24.V.69, 
Podoclavella maluccensts 
Stolonica australiy (aggregates } 
Polycarpa pedtunculata 
Pyura irreaularis 
Herdmania momus 
Microcosm squamiger 


2. on surface of rocks: slow current; depth 6m; 
24.¥.69. 
Leptoclinides reticulatns 
Phatlisia depressiuseula 
Axcidia svadneyeisis 
Srolenicu carnosa 
Polycarpa papillate 
Polycarpa pedaneulata 
Pyiiraaustralis 
Pyura irregularis 
Halocynthia hispida 
Microcesmius stolonifera 
Microeosmus squimiger 
Crenicella antipoda 


3. epizoic on Amphiholis .antarctica; moderate 
current. 2 m/sec; depth 12 m; 19.v.71. 
Ratryloides leachi 
Pyura australis 
Herdmania momus 


Orr Beacu Hut, 1 km off Port Vincent: on 
travertine; no wave action; slight current; depth 
4m; 24.11.69, 

Ascidia sydneyensis 

Pyura irreeularis 

Halocynthia hispida 

Microcosmus nichollst 

Microcosmus helleri 


ASCIDIANS OF SOUTH AUSTRALIA S| 


Oronres Rank, off Port Vincent: 20 mz 26.iii,66. 
Sycozea cerebriformis 


Tarrey SHOAL, St. Vincent Gulf: depth indicated 
for each species; Feb. 1969. 


1. Sluggish current, sandy bottam. 
Phatlusia depressiuscula (@m 
Ascidia sydneyensis 12m 
Palycarpa pedunculata’ 16m 
Cremidocarpi etherideli J2m 
Pyura scoreshiensis 16m 
Hylocynthia hispida 12m, tom 
Micracosmus hellerpi 12m 


ta 


Moderate current (lo Im/sec); travertine bot 
tom covered by shallow sand; depth indicated 
for each species, 

Aplidium colelloides \8m 

Palycarpa elavata 20m 
Polvearpa pedunenlata 
Pynra australis 20m 
Haloevuthia hispida 
Pyipra scareshiensis 


[8 m, 20m, 272 m 


22m 


3. Mostly sand with some: travertine outcrops; 
depth 23 im. 
Sycozoa cerebriformis (on rock) 
A plidium colelloides 
Polycarpa clavata 
Pyura yittata 


4, Strong current (to 2m/sec); sheet fraverline: 
depth 24 m, 
Polycitor gigémmeum 
A plidium pliciferum 
Polycarpa peduneulata 
Pyura aistralis 


Upper Sr. Yincent Gur: on sandy bottom in 
Pasidonia australis community; moderate current 
(1 1 m/see.); depth [0-11 my: 4.1.67. 

Leptoclinides hingi 

Pyura spinifera 

Prura irregularis 

Halocynthia hispida 

and growing on razor shell Pinna dolohrata: 

Sycozod cerebrifjermiy 

Aplidium rubricollium 

Ascidia gemmata 

Polycarpa pedunculata 

Herdmania momns 


Orr Port GAwLer, St. Vincent Gulf: growing on 
Pinna and on Celicpora spp; slow current; depth 
18-20 m; 11.71.67, 


Sycazoa cerebrifarmis 
Leptoclinides rufus 
Phallusia depressiuseula 
Ascidia gemimata 
Botryfloides nierum 
Polycarpa papillata 
Palycarpa pedunculuta 


Ore Ourer Harnovur, St. Vincent Gulf: on Pinna: 
slow current; depth 8 m; 2.xii.68. 


Halocynthia hispida 


Orr SkKMAPHORE, St. Vincent Gulf: in sparse Posi- 
donie community, silly bottom: slow current; 
depth 31 m; 274.69. 

Polvearpa pedunculata 

Pyura scoresbiensts 

Microcosmus squarmiger 


Orr Semapuore,, St. Vincent Gulf; silty bottom: 
slow current; depth 24 m: 28.xiLG8. 
Polycarpa pedunculata 


Ort GRANGE, St. Vincent Gulf: rocky bottom: 
slow current; depth 18 mt 7.xti.68. 

Phallusia depressinsenta 

Polxvcarpa pedunculata 


Ocr Grancr, St. Vincent Gulf; in Posidonia com- 
munity on shell; depth & my 7.x1i.68. 
Pyare irreeularix 


Orr West Beacu (uhont 3 km), St. Vincent Gult: 
on rocky bottom; depth 10 m; &.vi.68. 


Ascidia thempsant 
Boatrylinides magnicaecus 
Polyearpa peduncalata 
Cnemidocarpa etheridgii 
Haloeynthia hispida 

Pytira australis 

Pyura irregitlaris (agprepates ) 
Microcasrmus sqnamiger 
Micracosmus nichally¢ 


Orr West Beacu (about 7 km), St, Vincent Gulf: 
in Posidonia community, slow current, depth 12— 
20 m, 27-.xii.46, 

Endistoma pyriforme 

Phallusia depressiiuscula 

Batrxviloides nigrum 

Palycarpa pedunculata 

Pytra australis 

Herdmania momus 

Halocynthia hispida 


Orr West Bracu (about 9 km), St. Vincent Gulf: 
on silty bottom; slaw current; depth 20-25 mi: 
27.xiL. 46. 

Phallusia depressiuxenla 


Orr Broapway or GLENELG (several stations), 
St. Vincent Gulf: on sandy bottom: slow current; 
depths inditated for each species: 10,xi.68. 
Sycazad tennieaulis (on scallop shell; 22 m) 
Patycarpa pedunculata 616m 
Pyura australis 12m 
Halocynthia hispida & m 


Orr Gienrre (5 km), St. Vincent Gulf: rocky 
bottom; slaw current; depth 13 m: {3.v.67, 
Ascidia gemmata 
Polycarpa papillata 
Palycarpa peduneulata 
Terdmania momus 


Orr Grenenc (1.5 km), St. Vincent Gulf: on 
Posidania roots; depth 6 m: 30.Vv.70. 

Palycarpa pedinculata 

Pyura irrepularis 

Talocynthia hispida 

Microcosmus squamiger 


52 PATRICIA KOTT 


Orr GLENELG (14% km). St. Vineent Gulf: depth 
35 ms; 4,1x.69, 


Herdmania momus 


Orr SeAcuiFr, St. Vincent Gulf: in Posidonia 
community, on sandy bottom, fair sediment, slow 
current; depth 16 m: 21169. 


Ascidia aclara 


Orr Seacuer, St. Vincent Gulf: on Amphibolis 
autarctica, slow current; depth 9 m; 28.1x,68. 


Botrylloides nigrum—with sponge 


Orr Hatvet? Cove (3-5 km), St. Vincent Gulf: 
on silty bottom; slow current; depth 15-22 m; 
27,xii.46. 

Phalluyia depressiuscula 


Borryllus schlasseri 
Polycarpa pedunculata 
Pyura australis 

Pyura spinifera 


Orr Port Stanvac (6.4 km), St. Vincent Gulf: 
on steel wreckage (“The Barges”); slow current; 
depth 30 m; 26.11.66. 

Phaltlusia depressiuscula 

Halocynthia hispida 


Index to Genera and Species 


Page 
Aplidium colelloides » 15 
Aplidium pliciferum . 13 
Aplidium rubricollum So on Ul ODS 
Ascidia actara . . - ok 27 
Ascidia gemmata ate oh 26 
Ascidia sydneyensis Piotr orc 24 
Ascidia thompsoni . , 27 
Atapozoa fantasiana oa. gee: BF 
Botrylloides leachi ot . on, 228 
Botrylloides magnicoecum gs, 330 
Botrylloides nigrum 7, a coe iP 30 
Botryllus schlosseri - ie aie eM 
Clavelina baudinensis _ .. fA 
Cnemidocarpa etheridgii tg ert ee 
Corella eumyota oo... ... .. .- -. 23 
Ctenicella antipoda n.sp. ee AA 
Cystodytes dellechiujel . .., . - ai 
Didemnum candidum .. 19 
Didemnum lambitum eke he yee 2S 
Didemnum moseleyi rr 
Didemnum patulum Wont care LB 
Distaplia viridis reais ce eee A 
Echinoclinum verrilli oe ae Oe 
Eudistoma pyriforme , > & 9 
Eudistoma renieri - \ op tee ae WG 
Halocynthia hispida ar, _ Al 
Herdmania momus - ; 41 
Leptoctinides kingi ere saeli> adalat LY 
Leptoclinides reticulatus _. 18 


Orr YANKALILLA Bay, St, Vincent Gulf: in 
Amphiholiy community. sandy bottom: slight 
surge; depth as indicated: 18.11.67. 

Pyuraanustraliy 20m 

Halocynthia hispida 20m 

Crenicella antipoda 15m 

Page 

Leptaclinides rufus ; tie tam esp 
Microcosmus helleri 43 
Microcosmus nichollsi 42 
Microcosmus squamiger - - = 43 
Microcosmus stolonifera = ef “43 
Molgula mollis ; ; 45 
Oculinaria australis 29 
Phallusia depressiuscula ., 2... 4... 23 
Podoclavella cylindrica .- 1. te 
Podoclavella moluccensis : 5 
Polycarpa clavata . ee . 33 
Polycurpa papillata .. t Sie som 4 
Polycarpa pedunculata —- _f.. Pegs 
Polycilor giganteum tenet nie she 
Polysyneraton orbiculum - = 21 
Pseudodisioma cereum =... 4, 4.) «612 
Pyura australis = - et «se 
Pyura irregularis ee ee a nee 
Pyura scoresbicnsis 1.sp. a 36 
Pyura spinifera : , 39 
Pyura vittaia |) 4. -. + 
Rhodosoma turcicum . 23 
Ritterella herdmania — ee. Pk 
Stolonica australis _. He tte 
Stolonica carnosa a 28 
Sycozoa cerebriformis 
Sycozoa tenuicaulis... ny 
Synoicium papilliferum . ; oe ate | 16 


Trididemnum spiculatum ih tte ee DG 


NEW FORM SPECIES OF POLLEN FROM SOUTHERN AUSTRALIAN 
EARLY TERTIARY SEDIMENTS 


BY WAYNE K. HARRIS 


Summary 


Sixteen new form species of dispersed pollen grains; Sparganiaceaepollenites barungensis, 
Amosopollis dilwynesis, “Triorites” psilatus, Tricolporites valvatus, Triporopollenites gemmatus, 
Ericipites crassiexinus, Sapotaceoidaepollenites rotundus, Proteacidites confragosus, P. tripartitus, 
P. kopiensis, P. tortuosus, P. clintonesis, P. fromensis, P. varius, P. wilkatanaensis, and P. 
concretus and one new form genus, Gambierina, are described from early Tertiary sediments from 
southern Australia. 


NEW FORM SPECHFES OF POLLEN FROM SGUTHERN AUSTRALIAN EARLY 
TERTIARY SEDIMENTS 


by Wayne K. Harrts* 


Summary 


Sixteen new form species of dispersed pollen grains: 


Sparganiaceacpallenites hearungensin, 
i a 


Anwxopollis dilwynensis, “'Triorites" psilatus, Priculporites valvatus, Triperapollenitey cemmatus, Eri- 


cipites crassiexinus, Sapotacevidacpollenites 


kopiensis, P. tartuasus, P. elintonensis, P. fromensis, P. varius, P. wilkarandensis, 


rotundius, Proteacidites. confragasus, P. 


lripartitus, Py 
and #.. comerepirys 


and one new form genus, Garmhierina, are described from early Tertiary sediments from southern 


Australia, 


Introduction 


This paper describes several new species and 
anew genus, Gambrerina, that were mentioned 
4S Manuscript names by Harris (1971) in an 
account of the palynology of Tertiary sedi- 
menis in the Otway Basin. These forms were 
considered to have some biostratigraphic sig- 
nificance. 


Previous taxonomic studies of Tertiary 
apgiosperm pollen from Australia are limited 
ta the works of Coakson (1947, 1950, 1953, 
1954, 1957 & 1959), Cookson & Pike (1954) 
and Harris (196Sa), Dettmann & Playford 
(1968) described four new angiosperm pollen 
species from Upper Cretaceous sediments from 
eastern Australia and some of these probably 
extend into the carly Tertiary. 


The preparation technique is that outlined 
by Harris (1965a) und the descriptive termino- 
logy is largely adapted from Erdtman's glossary 
(1952). Dimensions are based on fifteen or 
more specimens. Biostratigraphic data are 
‘based in part on unpublished studies by the 
author and on Harris (1971) and McGowran, 
Lindsay & Harris (1971). Sample data are 
presented jn the appendix. 


All co-ordinates are from the Leitz Ortho- 
plan (715494) microscope in the Palynology 
Laboratory of the Geological Survey of South 
Australia and Holotypes (catglogue numbers 
prefixed Py) are deposited in the Geological 
Survey Palaeontological collection, 


Systematic Palynology 
Genus SPARGANIACEAEPOLLENITES 
Thiergart 1937 


Type species: Sparganiaceaepollenites polyga- 
nealis Thiergart 1937: 307. 


Sparganiaceaepollenites barungensis Sp. tov. 
FIGS. 1-3 


Pollen monoporate. sphucrojdal to slightly 
bilateral. Pore circular (3-4 pin diam.) with 
incrassate margin 1-1.5 ym wide. Exine 2 ~m 
thick, sexine as thick as nexine, reticulate. 
Reticulum undifferentiated over the grain, 
lumina 1-1.5 pm diam, Dimensions: Equa- 
torial diam. {8 (22) 25 ym. 


Holotype: Preparation and slide number— 

$T325/15, 42.1: 110.8. Py 195. Figs. 1, 2 

Type localify; Hd. Bartinga Bore 4 at 65.5 

m. Clinton Formation, 7Lower Miocene. 

Distribution: This species first occurs in the 

Upper Eocene and continues through to the 

Upper Tertiary. 

Comparison and affinity: The pollen figured 
by Couper (1960, pl. 9, figs. 21, 22) as Typha 
8p. appears to be very similar to §. barunvensis. 
S. harungensiy differs from Aglaoreidia Erdt- 
man in not having a differentiated reticulum. 
“Monoporitey” subreticulata Cookson has a 
wider rim to the pore, The specics here des- 
eribed appeared under this generic name in 
Harris (1971) and McGowran, Lindsay & 
Harris (1971). S: magnoides Krutzsch (1970) 


ST a ee 


* Geological Survey of South Australia, 169 Rundle Street, Adelaide, S.Aust. 5000 
Published with the permission of ihe Acting Director of Mines, 


Trans. R. Soc. S, Aust, 96, Part 1, 29 February, 1972, 


54 WAYNE K. HARRIS 


approaches S. barungensis in size but hax a 
wider meshed reticulum. S. batiurgensiv 1s 
very similar to pollen of Typha and. Sparge- 
niu, 


Genus AMOSOPOLLIS Cookson & Balme 
1962 


Type species: Amoxepollix cruciformis Cook- 
son & Bulme 1962: 97, 


Amosopollis dilwynensis sp, noy. 
FIGS. 4. 5 


Synonymy. Amosopollis erucifarmiis sensu 

Harris 19650: 97, Ph 29, fig, 26, 

Pollen grains in rhomboidal tetrads.  Indi- 
vidual grains prolate to sub-prolate, [xine 2 
am thick, psilate to seabrate and finely granu- 
Jale, except near the margins of the aperture 
where grana 1=1,5 jm diam. are present. Aper- 
ture ix a long gaping sulcus extending the full 
length of the grain, Margins of sulcus not 
ragged, Dimensions: (10 specinrens) Overall 
diam, of tetrad 50 (60) 68 pm. Individual 
arains 22 (34) 40 ,.m diam, 

Noloiype: Preparation and slide number— 

§T209/ 2, 39.3; 100.7. Py O15. Figs. 4. 5, 

Type locality: Ditwyn Bay, Victorian, Pebble 

Point Formation, Paleocene. 


Distribution; A. dilwynensis is a rare specics 
hut has been observed in Paleocene sedi- 
ments fram the Murray and Otway Basins, 
and a Similar forny hus been reported (Harris 
1965h) from Queensland in sediments of 
similar age. 


Comparixon and affinity: A. dilwynensis is 
in general Jarger than the genotype but can also 
be distinguished by the psilate-scabrate sculp- 
tre und more importantly by the straight mar- 
fins of the sulcus. 


Genus TRIORITES Cookson ex Couper 1953 
Type species (hy subsequent designation of 
Couper 1953, p. 60): Triorites magnificus 
Cookson 1950, 
‘“Triorites” psilatus sp. nov, 
FIGS. 6, 7 
Pollen radiosymmetric, isopalar, oblate, 
triorate. Amb sub-triangular. sides straight to 
slightly convex. Ora sunken, 2-4 pm wide, 
circular Exine 2m except around apertures 
where it thickens to 3 or 4 pm. Enxine psilale- 
Dimensians: Equatorial diam, 24 (32) 40 pm. 


Holotype; Preparation and slide number— 
5964/1. 32.8: 100.6. Py 411. Fig. 6. 


Type locality: Polda No. | Bore at 55.1 m. 
Poelpena Formation, Middle Bocene. 
Distribution: ‘Vhis species is a very common 
form throughout the Lower Tertiary in 
southern Australia. ft first appears in the 
Princetown Member and ranges through to 
ihe Lower Miocene. The upper limil has 
nor been determined, 


Camparivon and affinity. “T" psilaris is com- 
parable and may be conspecific with "7." 
yewbratas Couper, "The ornament on the latter 
however is scubyate. This species would more 
appropriately be placed in a new genus (see 
section on Trieritey below). 


Genus VRICOLPORITES Cookson 1947 
Type species: Tricolporites sphacrica Cookson 
1947; 195; genus monotypic when proposed, 
Tricolporites valyamms sp. nov, 

FIGS, &, 9 

Pollen radiosymmetric, prolate tricolpurpie 
Amb in equatorial view ellipsoidal. Apertures 
compound, colpi reaching to within 3 or 4 jn) 
of the poles, margins strongly invaginated to 
about § pm. Equatorial aperture orate, 5-8 
pm diam, Kxine 2-3 am thick unornamented. 
Dimensiony: Pole diam. 45 (52) 55 jum, eque- 
torial diam, 30 (35) 39am. 

Holoiype: Preparation and slide nurmber— 

ST241/12, 35.1; 98.1. Py 176, Fig. 9, 


Type docality: Lake Torrens Bore 3A at 


247.8 of “Wilkatana Formation”, Middle 
Focene. 

Distribution: Common in the “Wilkatana 
Formation" but less common in other 


Middle Kocene (Proleacidites confragosus 
Zonule) assemblages, 


Camparison und affinity: The strongly. invagi- 
nated colpi and psilate exine make this a very 
distinctive species, Its natural affinilies are un- 
known, 


Genus ERICIPITES Wodehouse 1933 


Type species: Fricipites longisitlearis Wode- 
house 1933: 517. 


Ericipites crassiexinus sp. nov. 
FIGS. 15, 16 


Pollen united in tetrads. Individual grains 
indistinctly tricolporate, tetrahedral in shape 


FARLY 


and strongly united in the tetrad. Exine 2.5- 
3.5 ,m thick, sexine as thick as nexine psilate. 
Aperttires complex, colpi about 14 pm long 
and 1.5 pm wide, Pores indistinct and difficult 
ta detect, 2 wm diam, Dimensions: Overall 
diam, 35 (42) 33 am. Individual grains 24 
(3 35 pon diam 


Holarype, Preparation and slide mumber—- 
S660/ 1, 52.9: 96.9. Py 415, Fig, 15. 


Type locality: Bore, Hd, Cummins at 35,7- 
43.30 om. Wanilly Formation, Middle 
Eocene. 


Distribution: Often wu yery coramon. form in 
middle and upper Eocene sediments. 


Compariven and affinity: The psilate nature of 
the exine and the larger size of this species cis- 
tinguishes it from A. scabrases Haris. Pollen 
of this type characterise the Order Ericales. 


Genus TRIPOROPOLLENITES (Pflug) 
Thomsen & Pfluz 1953 
Type species! = Triperopollenites 
Pflug in Thoms. & Pf, 1953: 84. 
Triporopollenites gemmatus sp. nov. 
FIGS, [0, 11, 13, 14 

Pollen occasionally free but most commonly 
united in tetrads. Tetrads 34-40 2m in overall 
diam. Individual = pollen = radiosymmetric, 
oblate, sub-isopolar, triorate. Amb sub-trian- 
gular with straight to convex sides. xine 4—5 
am thick (including ormament), Sexine and 
nexine difficult to separate but nexine appears 
to be thicker than ‘sexine. Exine covered with 
verrucae 2-3 «wm wide, sphacrical and 2 ym 
high. Verrucue separated from each other (by 
2-3 am) by granulate ornament. Apermres 
obscured by ornament, porate or orate open- 
ing 1.5-2.5 pm wide, Dimensions; Individual 
pollen, equatorial diam, 25 (29) 31 jam. 


Holotype; Preparation and slide number— 
$547/1, 31.7: 98.4, Py 720. Fig. 11. 


Type locality: Lake Cootabarlow Bore 2 at 
163.4 m. Great Artesian Basin. Murn- 
peowie Formation, Upper Eocene. 


Disrriburion:; Appears to be testricied to 
Middle and middle-upper Eocene sediments 
from the Pirie-Torrens and Great Artesian 
Basins and Eyre Peninsula, 
Comparison and affinity; T. 2zemmatus is simi- 
lar to 7. bullis Gruas-Cavagnetto (1966) from 
the Sparnucian of the Paris Basin but this 


coryloides 


JERTIARY POLLEN SPECIES 55 


species is more of less circular and appears to 
have a more strongly thickened sim to the 
aperture: 


CGienus GAMBIERINA gen, nov. 
Type species: Triorites edwardsit Cooksoly & 
Pike (in part) 1954: 214, pl. 2, figs, TOT, 105, 
106, 


Diagnosis; Pollen raciosymmetric, oblate, 
lohate. angulaperturate, triorate. Apertures 
sunken. Sexine imperforate tectate, thinner 
than nexine, the two separated by a faintly dis- 
cernable baculate layer, which forms a “nick” 
point in the apertural region, Aperture fortyed 
by sexine larger than that of the nexine. Nexine 
thickens more rapidly than sexine about the 
apertures. Exine psilate. 


Figured specimen: Fig. 12, 


Remarks: The characters of the exine, the 
apertures and general shape distinguish this 
venus from Triorites. As Dettmann & Play- 
ford (1968, p. 86) have pointed out, the 
species figured by Cookson & Pike (1954, par- 
ticularly Ags, 104 and 105) as 7, edwardsii is 
distinct in being unthickened about the aper- 
tures, 


Dettmann & Playford (1968) summarised 
the present status of ahe genus Triarites but 
chose to continue using the diagnosis of Couper 
(1953) pending a review by the present author. 


Potonié (1960) clearly indicated that the 
two species T. mugniftcuy Cookson and T. 
efavatus Cookson were morphologically com- 
parable and distinct from other torms allocated 
to the genus. However, Potonié gave no indi- 
cation as ta where these other forms should be 
placed. 


It ig clear that T. magnificus and T. clavatuy 
are very closely related morphologically and 
perhaps phylogenetically, Indeed Cookson 
(1957, p. 49) voes so far as to state thar “there 
is little or no doubt that they were produced 
by closely related plants. Both species have 
the same shape, type of ora and exine stratifi- 
cation, and structure...“ Thus these two 
species form a natural grouping and all other 
species assigned to the genus ate better acco- 
modated elsewhere. Couper’s (1953) diagnosis 
is too broad and suggestive of a suprageneric 
category. Mildenhal) & Harris (1971) have 
reached similiar conclusions. 


Genus SAPOTACEOIDAEPOLLENITES 
Pot. Thoms, & Thierg. 1950 


Type species: Saparaceoidacpollenites. (al. Pal- 
lemlex} vratttfesnes (Potonié) 1931: 3. 
Sapataceoidaepollenites rutundus sp. nay. 
FIGS. 17, 18 

Pollen radiosymmetric, subsphaeroidul to 
sub-prolate, four and less frequently three 
apertures. Apertures compound. Colpi 2/3 
length of polar axis, 2-3 ym wide. Equatorial 
aperture more or less circular, S=6 pm diam. 
and slightly elongate 0 an equatorial direction, 
Apertural margin prominently rimmed ancl 
thickened, xine 2-2.5 pm_ thick, nexine 
ubout as thick as sexing, Sesine psilate to 
finely scabrate. Dimensions: Polar diam, 30 
(36) 39 am, equatorial diam. 26 (33) 35 pm, 


Holotype: Preparation and slide number 
ST241/3, 45.7: 102.8. Py 167. Fig. (7. 
Type locality: Lake Torrens Bore 3A ut 
247.8 om. “Wilkatuna Formation, Middle 
Focene. 
Distribution: The species Orst appears in the. 
Middle Hocene and continues on Into the 
mid-Tertiary. 
Comparison and affinity: The closest resem- 
blance of this species is with Tricalperopollen- 
ites latizonatuy MeIntyre 1968, which is most 
commonly 3-aperturale, has a longer polar/ 
equatorial axis ratio and has a broad thickened 
zone of the exine in the equutorial region, The 
species ts very similar to pollen of the Sapo- 
taceae, 


Genus PROTEACIDITES Cookson 
ex Couper 1953 


Type species: Proteacidites adenamhotdes 
Cookson (950: 172, designated by Couper 
1953; 42, 


Remarks, The genus Proteaciditey wecomodales 
ul present. a wide variety of forms described 
fram both the Southern and Northern Hemis- 
pheres. Some from the latter clearly do not 
belong in this genus but until a review of the 
Australian forms by the author is complete 
(and on present evidence the genus will be split 
into three or more genera) comment on these 
is reserved. Although the following new spe- 
cics show a wide variation in form with regard 
to aperture construction and exine stratification 
and ornamentation they ‘will be described 
under this genus but will be further reviewed 
in a forthcoming paper. 


56 WAYNE K. ITARRIS 


Proteaciites coufragosus sp. nov. 
FIGS, 19-22 


Pollen sub-isopolur anguluperturate, oblate. 
tiporate. Arub triangular with slightly conves 
sides. Pores simple subcireular 6-7 pm diam.. 
obscure. Exine 4-5 pm thick, Sexine three 
limes us thick as nexine, heavily ornamented 
with a dense reticulum, lumina 3-4 pm diam.- 
polyvonal wind made up of single rows of dis- 
tinct bucula 1-15 um diam. Dimenstons: 
Eyuatorial dian, 34 (60) 69 pm, 


Halorype: Preparation and slide number— 
Sr24t/9, 41.4: 105.7, Py 173, Figs. 19, 


21, 22. 


Type locality: Lake Torrens Bore 3A. aol 
2478 m “Wilkaturm Pormation’. Middle 
Eocene. 


Distribution: An index form for Middle 
Eocene sediments, PF, confrugosus has been 
recorded from the North Maslin Sands, the 
Renmark, Poelpenu and Wanilla Formations 
and the Burrungule Member of the Knight 
Formation, 


Comparivan and affinity: This is 4 striking spe- 
cies and is ¢learly distinct from any other 
known in the genus. 


Proteacidites tripartitus sp. nov, 
FIGGS. 23 25 

Pollen sub-isopolar, oblate, angulaperturate, 
triporate, Amb triungular with more or less 
straight sides, Apertures sub-circular, simple 
but obscure 2-2.5 ym wide. Exine 2.5-3 pm 
thick, Sexine half as thick as nexine, fove- 
alate, Lumina ca. 1 am diam., slightly smaller 
at the poles and towards the apertives. Muri 
2-3 ym wide. Nexine thickens. to 5 ym at 10 
pm from the apertures. Pore “canal” 7-8 yam 
long. Déinensions: Equatorial diam, 27 (30) 
34 pitt, 

Holoi\pe: Preparation and slide nuniber — 

$650/1, 32.2: 99.8. Py 406. Pigs. 24, 25 

Type locality’ Hd, Cummins Bore at 114- 

116.4 im. Wanilla Formation, Middle 

Eocene, 


Disiribution: The species first appears very 
high in the Princetown Member of the Dil- 
wyn Formation but does not became com- 
mon unfil the Miidle Eocene. 


Compurixon and affiniry: The detail of the 
apertures closely resembles that found in P. 
latrohensts Harris and P. concretus but is dis: 
linguished bY the characteristic ornament, 


EARLY TERTIARY 


Proteacidites kopiemsis sp, nov, 
FIGS. 26, 27 

Pollen sub-sopolar, oblate. angulaperturate, 
Iriporate, Amb triangular, sides straight or 
nearly so. Apertures subcircular. simpic. 7-8 
pity diam. Exine 2m thick and slightly thicker 
in the equatorial inter-aperturate regions. 
Sexinc about half as thick as nexine, orma- 
mented with a reticulum, Muri 1-15 pm 
wide, Lumitta 2-3 am diam, at the equator 
and decreasing gradually to 1 um towards the 
apertures and polar regions.  Dwnenstons: 
Equatorial diam, 36 (40) 47 ym, 


Hololype: Preparation and slide number~- 
S560/1, 26.1: 106.9. Py 393. Fig, 26, 


Fiipe tocality: Pala No. | Bore at 37,5 m, 
Poelpena Formation. Middle Eocene. 


Distribution: The species is present in the 
uppermost section of the Princetown Mem- 
ber of the Dilwyn Formation and continues 
in all basins into the middle-upper Eocene. 
IL does not appear to range higher than the 
Triorites magnificuy Zonule, 

Comparison and affinity: This species is teaditv 

Wistinguished from other Proteacidires spp. by 

the characteristic ornament pattern. 


Proteacidites tortuosus sp, nov, 
FIGS, 28. 29 


Pollen sub-tsopotar. oblate, anguluperturate. 
triporate. Amb rounded triangular, sides con- 
vex. Pores simple 13-15 »zm wide. Exine 5 
pm thick. Nexine thicker than sexine. Sexine 
ornamented with scattered verrucae, 2 «um 
wide and up to 6 xm long, rounded in optical 
section and 2 ym high. Areas between these 
elements psilute. Dimensions: Equatorial diam. 
53 (55) 58 pm 


Holorype: Preparation and slide number— 
$563/2, 32,2; 99.8, Py 409. Figs, 28, 29, 
Type locality: Polda No, | Bore at 55.1 mm. 
Poelpena Formation, Middle Eacene, 


Distritiaion: This species has heen recorded 
from Middle Eocene sediments on Eyre 
Peninstila, Poelpena and Wanilla Porma- 
tions. 


Comparison and affinity: The large distinctive 
verrucae, thick exine and rounded sub-triangu- 
lar shape separate this species from other spe- 
cies described here. P, tortuasus differs fram 
P. nibereudatuy Cookson in being smaller. The 
vermicae are not arranged in u reticuloid pat- 
tern and are not confined to a sphencal shape 


POLLEN SPECIES 57 


Proteacitites clintonensis sp. nov. 
FIGS. 30-34 
Pollen sub-isopolar, oblate angulaperiurate. 
triporate, Awb more or less triangular with 
eoncave sides, Pores circular 2435S ym in 
diam, Exine 3 jm thick, sexine slightly thinner 
thin nexine. Capita of bacula coalesce to 
form groups Up to 7am wide and shmw an LO 
pattern, Elements rounded in optical section, 
Nexine in region of pores, alternately thick and 
thin, The sexine is readily lost by corrosion, 
Dimensions: Equatorial diam. 62 (75) 98 yum. 
Hoelompe; Preparation and slide aumber— 
$705/1, 31,3: 105.2. Py 405. Figs, 32-34, 
Type fecaliry: Poyniz Bore, Hel. Ettrick at 
94.5 om. Renmark Beds, middle-upper 
Eocene. 
Distribution: The species is almost ubiqui- 
tous in Eocene sediments and is particularly 
common in the Priarites staynificuy Zonule, 
{| ranges from Middle Eocene to at feast 
Lower Miocene, 
Comparison and ajfinity: This species is similar 
to FP. rectomerginis Cookson but has much 
larger apertures and strongly concave sides. 
Figure 30 more closely resembles P. reeto- 
mmarginus with its finer ornament, larger size 
and straighter sides. It is possible that the two 
forms intergrade, Cookson’s figure (1950. fig. 
27) of P. rectomarginuy appears ta show some 
thickening of the nexine about the apertures. 
The species is distinguished from. P. jncurvelius 
by the nature of the sculpture and the charac- 
teristic aperture. The exine dacs not rhin 
markedly near the apertures as it does in P, 
irearvatus, 


Proteacidites fromensis sp, nov. 
FIGS. 35-38 

Pollen sub-tsopolar, oblate, angulaperturate. 
triporate, Amb triangular, sides strongly con- 
cave, Pores simple, circular, 5 ym in diam. 
Exine 2,3-3 »m thick. Sexine slightly thinner 
than nexine in the inter-angles and thins to- 
wards the angles, evenly granulate to scahrate. 
Nexine thickest im the inter-angles, Désrten- 
sions: Equatorial diam. 61 (65) 70 um, 

flolotype: Preparation and slide number 

S17/2. 35.1: 1014, Py 408. Figs, 35-37. 

Type locality; Lake Eyre Bore 20.at 73.2 m. 

Murnpeowie Formation, Paleacene. 


Distribution: P. fremensis is restricted to and 
characteristic of Palacocene sediments and 
is most cormmmon in the Murray and Great 
Artesian Basins, 


38 WAYNE K. HARRIS 


Comparison and affinity: The strongly concave 
sides of the amb, size, and the scabrate orna- 
ment separate this species. fram others in the 
genus, It differs from P. granoraius Couper 
ive that the ornament does not become coarser 
around the apertural region. 


Proteacidites varius sp. nov. 


FIGS. 39-42 


Pollen small, sub-isopolar, peroblale, angu- 
japerturate. Amb triangular with straight or 
slightly Concave sides. Apertura] pores, three, 
2.5 »m diam. Exine 2-2.5 pm thick. Sexine 
much thinner than nexine and thins markedly 
near the apertures, Necxine thins toward the 
apertures with loss of Jendonexine elements 
near ihe wperture. Ornament 0.5 pm high, 
consisting of fused groups of bacula ca. 1 pm 
djam, Groups becoming smaller to absent 
near apettures. Dimensivns: Equatorial diam. 
20 (25) 37 am, 


Holaryye: Preparation and slide number— 
§705/1, 37.2: 111.0. Py 399, Figs. 39, 40. 


Type locality: Poyntz Bore, Hd. Ettrick ut 
94.9 m. Renmark Beds, Upper Eocene. 


Distribution: A common species in middle 
and upper Eocene assemblages in the Murn- 
peowie and Poelpena Formations in parti- 
cular, 


Comparison and affiniry; Vhe nature of the 
ornament distinguishes the species from P. 
reliculainsy Cookson and P. xymphyonemoides 
Cookson, The churucteristic nexine structure 
(see particularly Fig, 41) is distinctive, The 
relitionship of this species. lo P. obscurus 
Cookson is not clear. Her figured specimens 
(Cookson 1952, figs. 30, 31) have lost most 
of the sexine, The specics described here 
shows thinning of the exine about the apertures 
rather than slight thickening and is not “lanect- 
shaped” as described for P. ebscurus. 


Proteacidites wilkatanaensis sp, nov. 
FIGS, 43-47 
Pollen sub-isopolar, oblate, angulaperturate, 
triporale. Amb triangular with straight lo con- 
cave sides slightly bulging about [0 ~m from 
wpertures. Apertures circular 4-6 pnt diam, 


Exine 4 um thick, thinning rapidly near the 
Apertures. Nexine about 3 times as thick as 
sexine, thinning and apparently losing the basal 
layer near the apertutes, Sexine consists of a 
thin baculate layer and an ectosexinous layer 
formed of united baculate elements. giving a 
fuw rugulate ornament. Rugulae L-3 ,«m 
long, lesa than 1 pm wide. Dimensions: Equa- 
torial diam. 51 (55) 67 2m. 


Holotype. Preparation and slide number— 
$2273/'4, 44.1: 106.7. Py 720. Figs, 43. 
44. 


Type locality: Bore near Ediacara at 280,4— 
283.5 m, “Wilkatana” Formation, middle- 
uppet Eocene. 


Pistribuion: The species is commonly ob- 
served in miiddle-upper Eocene sediments in 
most basins. 


Comparison and affinity: The species differs 
from P. ineurvans Cookson and P. clintonensis 
in not being puhcti-tegillate. The ornament is 
similar to that of P, varius bul the species is 
much larger and does not show the charac- 
teristic structure of the nexine wround the aper- 
tures, 


Proteacidites concretus sp. nov. 
FIGS, 48, 49 


Pollen sub-isopolar. oblate, ungulaperturate, 
Iripotute. Amb triangular with straight sides, 
Apertures circular 1.5 ,m diam, Exine 2 um 
thick but thickens in the region of the aperture 
to.4 am and forms a pore “canal” 5 pm long. 
Exine faintly and evenly scabrate to finely 
spinulate. LO pattern distinct. Dimensions: 
Equatorial diam. 25 (28} 32 ym. 


Holatype: Preparation and slide number— 
$360/2, 35.8: 105.5. Py 404. Fig. 48. 


Type locality: Kopi Anomaly KR9 at GT om. 
Peolpena Formation, Middle Eocene. 


Distribution: A common species in most 
Locenge sediments, 


Comparison und affinity: This. species is most 
closely similar to P, farrobensis Harris, parti- 
colacly in the nature of the aperture. It differs, 
however, from this. species by the nature of the 
ornament. (P. farrabensis has a scrobiculate 
pattern.) 


EARLY TERTIARY POLLEN SPECIES 59 


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Sporen). Geol. Jb, 65. 35-70. 

THIERGART, F. (1937).—Die Pollenfora der 
Niedetlausitzer Braunkohle, besondets im 
Profil der Grube Marga bei Senftenberg. Jh- 
Preis. geol. Landesanst. BereA kad. 58, 222- 

S56. 

Tuomson, P. W., & Prruc, H. (1953).—Pollen 
und sporen des mittelcuropiischen Tertiirs, 
Palacontographica B, 94, 1-138. 

Wooexouse, R. P. (1933),—Tertiary pollen JI. 
The oil shales of the Eocene Green River 
ternation, Bull, Torrey Bot. Club, 60. 479- 
524, 


60 


WAYNE K. HARRIS 


APPENDIX 
SAMPLE DATA 


Depth in metres Type of Sample 
Bore name (or outcrop) (feet in parenthesis) Formation Basin Lovality Sample No, 
Nullabor No. 6 105-122 (344-400) Pidinya Eucla Lat. 31°09'00"S Percussion $623 
Long. 131°12’30"E sludge 
Polda No. | 37.5 (123) Poelpena Polda Lat. 33°33’00"S Core SSal? 
Long. 135°20’00”E 
Polda No, t 51.8 (170) Poelpena Polda Lat. 33°33/00"S Core $562 
Long, 135°20’00”E 
Polda No. | 55.1 (ABU) Poelpena Polda Lat. 33°33/00"5S Core 5863 
Long. 135°20'00 TE 
Polda No. | 57.9 190) Poelpena Poida Lat. 33°33/00"S Core S564 
Lone. 135°20’00” E 
Kopi Anomaly K.R.9 61 (200) Poclpena Unnamed Lat. 33°24'10"S Care S360 
Long, 135°44’45"E 
Hd. Cummins (W.Con.Res. 35,7-43.3 (117-142) Woanilla Cummins Lat. 34°15710"S Percussion $660 
adj. Sec. 16) Long. 135°40’45"E sludge 
Hd, Cummins (W,Con.Res. 114-116,.4 (374-382) Wanilla Cummins Lat. 34°15°10"S Percussion 5650 
adj, Sec. 14) Long. 135°40°45”"E sludge 
Cummins school residence 32-39 (115-128) Wanilla Cummins Lat. 34°15/50"S Percussion S74! 
Long. 135°43'20"E sludge 
Lake Torrens Bore 3A 247.8 (813) “Wilkatana” Piric- Lat. 31°14’00"S Core S241 
Torrens Long. 138°01’45’"E 
Near Ediacara 280,4-283,5 (920-930) ‘'Wilkatana’™’ Piric- Lat. 30°48’34"S Cuttings $2273 
Torrens Long. 138°07/30"E 
Hd. Barunga Bore 4 65.5 (215) Clinton St, Vincent Lat. 33°45'°55""'S Core $325 
Long. 138°13'35”"E 
Lake Cootabarlow Bore 2 163.4 (536) Murneowic Great Lat. 30°16730"S Care 8547 
Artesian Long. 140°08'30” E 
E. A. Rudd Bore § 116.1 (381) Murneowic Great Lat. 31°13’00"S Core $1986 
Artesian Long. 139°52/50"E 
Lake Eyre Bore 20 73,2 (240) Murneowic Great Lat, 28°48’00"S Core SIT 
Artesian Long. 137°30’20”E 
Payntz Bore, Hd. Ettrick 94.5 G10) Renmark Beds Murray Lat. 35°O00°307"S Percussion $705 
Long, 139°31’45”E sludge 
S.E. side of Dilwyn Bay 1.8 m ahove buse Pebble Point Otway Lat. 38°44/00”S Gutcrop S208 
of formation Long. 143°10°30"E 
SE. side of Dilwyn Bay 1.2 m above base Pebble Point Otway Lai, 38°44/00"S Onterop S209 


of formation 


Long. 143° 10730" E 


EARLY TERTIARY POLLEN SPECIES 61 


Note: Unless otherwise specified the figures are X500 in normal transmitted light. NDIC refers to 
Nomarski Differential Interference Contrast. 


Figs. 


Figs. 


Figs. 
Figs 


Figs. 


Fig. 


Figs. 
Figs. 
Figs. 
Figs. 


Figs. 


Figs. 


Figs. 
Figs. 3 


Figs. 


Fig. 
Figs 
Figs 


Figs 


1-3. 


4,5. 
3 06,. 7. 
. 8,9. 
10, 11. 


12 


Figs. 1-12 
Sparganiaceaepollenites barungensis sp. nov., X 1250, Figs. 1, 2.—ST 325/15, 42.4: 
110.3. Fig. 1.—High focus. Fig. 2.—Mid focus. Fig. 3.—S 325/3, 39.7: 96.8. Pore 
in N.W. quadrant. 
Amosopollis dilwynensis sp. nov. Py 015, 38.7: 100.1. Fig. 5, X 1250. Notice the 
granulate margin of the sulcus. 
“Triorites” psilatus sp. nov. X 1250. Fig. 6.—S564/1, 32.8: 100.6. Fig. 7.—S562/1, 
37.7: 104.9. 
Tricolporites valvatus sp. nov. Fig. 8—ST 241/12, 39.0: 98.2. Fig. 9—Py 176, 35.1: 
98.1, X 1250, NDIC. 
Triporopollenites_gemmatus sp. nov. Fig. 10.—S650/1, 43.4: 104.5, single grain. Fig. 
11.—S547/1, 31.7: 98.4, tetrad. 
Gambierina edwardsii (Cookson & Pike) Harris comb. nov. ST 208/2, 32.1: 103.7. 


SSS 


Figs. 13-25 


T. gemmatus sp. nov. Fig. 13.—S547/1, 31.7: 98.4, X 1250 high focus. Fig. 14.— 
S741/2, 38.6: 106.8. 

Ericipites crassiexinus sp. nov. Fig. 15.—S660/1, 52.9: 96.9. Fig. 16.—S560/1, 19.2: 
107.2. 

Sapotaceoidaepollenites rotundus sp. nov. Fig. 17.—Py 167, 45.7: 102.8, X 1250, NDIC. 
Fig. 18.—S564/1, 29.5: 106.4. 

Proteacidites confragosus sp. nov. Figs. 19, 21, 22.—ST 241/9, 41.4: 105.7. Fig. 21, 
22, X 1250. Fig. 9 focused on ornament, fig. 10 focused on apertural region. Fig. 20, 
—ST 241/4, 35.6: 104.7. 

Proteacidites tripartitus sp. nov. X 1250. Fig. 23.—S560/1, 27.7: 100.8, NDIC. Figs. 
24, 25.—S650/1, 32.2: 99.8, sectional and high focus respectively. 


SSS 


Figs. 26-37 


Proteacidites kopiensis sp. nov. Fig. 26.—SS60/1, 26.1: 106.9, X 1250, NDIC. Fig. 
27.—S623/1, 16.6: 109.3, NDIC. 
Proteacidites tortuosus sp. nov. $563/2, 32.2: 99.8. Median and high focus respectively. 


Proteacidites clintonensis sp. nov. Figs. 30, 31.—S741/2, 47.8: 96.3. Fig. 31, X 1250, 


high focus on polar region. Figs. 32-34.—S705/1, 31.3: 105.2. Fig. 33, X 1250, high 
focus on polar region; fig. 34, focus on apertural region. 

Proteacidites fromensis sp. nov. §17/2, 35.1: 101.4. Figs. 36, 37, X 1250, Fig. 36, 
high focus on polar region; fig. 37, median focus on interapertural region. 


eee 


Figs. 38—49 
Proteacidites fromensis sp. nov. §1986/2, 27.0: 99.3. 
Proteacidites varius sp. nov. X 1250. Figs. 39, 40.—S705/1, 37.2: 111.0. High and mid 
focus respectively. Fig. 41.—S547/1, 22.7: 96.0. Fig. 42.—S705/1, 98.8: 44.3. 
Proteacidites wilkatanaensis sp. nov. Figs. 43, 44.—S2273/4, 44.1: 106.7. Mid and high 
focus respectively. Fig. 45—S705/2, 22.8: 105.9. Figs. 46, 47.—S2273/3, 33.2: 107.1. 
High and mid focus respectively. 
Proteacidites concretus sp. nov. X 1250. Fig. 48.—S360/2, 35.8: 105.5. Fig. 49.— 
$705/3, 26.9: 105.5. 


WAYNE K. HARRIS 


EARLY TERTIARY POLLEN SPECIES 63 


f4 


WAYNE K. HARRIS 


EARLY TERTIARY POLLEN SPECIES 6 


an 


VOL 96, PART 2 31 MAY, 1972 


TRANSACTIONS OF THE 


ROYAL SOCIETY 
OF SOUTH AUSTRALIA 


INCORPORATED 


CONTENTS 


Preiss, W. V. The systematics of South Australian Precambrian and Cambrian 


Stromatolites. Part 1 - - - - - - - - 67 
Mawson, Patricia M. The nematode genus Maxvachonia (Oxyurata: Cosmo- 

cercidae) in Australian reptiles and frogs - - - - 101 
Mawson, Patricia M. Three new species of the genus Cloacina Linstow (Nema- 

toda: Strongylata) from macropod marsupials - - - 109 
Schodde, R., Mason, I., & Wolfe, T. O. Further records of the Pitted-shelled 

Turtle (Carettochelys insculpta) from Australia - - - 115 


Wollaston, Elise M. The morphology and relationships of Muellerena wattsii 
(Harvey) Schmitz (Ceramiaceae: Rhodophyta) . - - 119 


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


THE SYSTEMATICS OF SOUTH AUSTRALIAN PRECAMBRIAN AND 
CAMBRIAN STROMATOLITES. PART 1 


BY W. V. PREISS 


Summary 


The methods of field study and detailed morphological analysis using three-dimentional 
reconstructions and thin sections, developed by one Russian school, were applied to the abundant 
Precambrian and Cambrian stromatolites of the Adelaide Geosyncline. Although other schools 
either demand formal taxonomy for algal remains only, or use informal descriptive nomenclature of 
morphologies which they believe are determined entirely by environment, it is concluded that valid 
and consistent stromatolite form-taxa can be distinguished by these studies. The recognition of 
stratigraphically restricted taxa suggests biostratigraphic subdivision and intercontinental 
correlations. 

New forms Acaciella angepena. A. augusta, Baicalia burra and Boxonia melrosa, and an 
indeterminate form of Acaciella, are described. 


THE SYSTEMATICS OF SOUTH AUSTRALIAN PRECAMBRIAN AND CAMBRIAN 
STROMATOLITES. PART L 


by W. V. Preiss“ 


Summary 


The methods of field study snd detailed morphological analysis using threc-dimentional secon- 
structions anid (hin secliuns, developed by one Russian school, were applied to the abundant Precambrian 
and Cambrian stromatolites of the Adeloide Geosyncline. Although other schools either demand fortnal 
taxonomy for algal remains only, or use informal descriptive nomenclature of morphologics which 
they believe are determined entirely by environment, it is concluded that valid and consistent atroma- 
tolitc form-taxa. can be distinguished by these studies. The recognition of Stratigraphically restricted 


taxa suggests biostratigraphic subdivision and intercontinental correlations. 
New forms Acacielia angepena, A. augusta, Baicalia burra and Roxenia melrosa, and an indeter- 


mitiate form of Acaciella, are described. 


Introduction 


Stromatolites are laminated structures 
formed in sediments, mostly carbonates, by the 
tapping and precipitation of sediment by mats 
of algae and bacteria. They are known through. 
out the sedimentary record, and are particu- 
larly abundant in otherwise unfossiliferous Pre- 
cambrian sequences. This observed abundance 
and the diversity of forms have made stroma- 
tolites potentially useful as index fossils, pro- 
vided that taxa can be defined which have stra- 
lgraphically restricted time-ranges, A group of 
Russian stromatolite specialists has been en- 
gaged in the systematic description and elassifi- 
cation of stromatolites for the past fifteen vears 
and their results stimulated this study of South 
Australian stromatolites in an attempt to apply 
biastratigraphic methods to the problems of the 
age and correlatron of the Precambrian 
sequence in the Adelaide Geosyncline. 


This paper is hased on the systematics sec- 
tion of a thesis submitted for the degree of 
Ph.D., University of Adelaide, It is necessary 
here to briefly discuss the taxonomy of stroma- 
tolites and particularly to formalize several new 
tixa meeded for subsequent discussions of bio- 
stratigraphy and palacoccology. This paper. the 
first. of three, will include yn outline of pre- 
vious studies, consideration of some taxonomic 
pringiples and problems, and descriptions of 
four new forms and one indeterminate form. 
The other parts will comprise descriptions of 
further forms and a discussion of the strati- 
graphic distribution of stromatolites. Stroma- 


tolite forms will be described in alphabetical 
order, 
Background 

This history of the early study of strama- 
toliles Was comprehensively reviewed by 
Masloy (1960), Although most researchers 
prior to 1914 sought an animal origin for these 
structures, for example Hall's Cryptezoon 
(1883) and Steinmann’s Gyrmrosalen (1911), 
Walcott'’s (1914) discovery of filamentous 
microfogsils in Precambrian stzomatolites from 
the Belt Series of Montans paved the way for 
the understanding of stromutolite formation by 
algae. 

Later workers clarified the role played by the 
algac. In particular, Black (1933) established 
that the algal mats of the Bahamas are poly- 
specific and that the mucilaginous filaments of 
the blue-green algae present trap detrital grains. 
Pia (1926) recognized the rock-building pro- 
perties of modern blue-green algac. Algal fila- 
ments were also found by Bradicy (1929) to 
aceur in stromatalites of the Eocene Green 
River Formation of Wyoming, 

In Australia, Mawson (1925) recognized 
stromatulites in the Flinders Ranges, and 
started a collection which was partly used in 
this study. During the 1930's fossil stromato- 
lites were described by numerous authors, the 
most. important being Young (1933a, (933b, 
1935), Fenton & Fenton (1931, 1933, 1936, 
1937, 1939), Johnson (1937, 1940) and 
Maslov (19374, 1937b, 1938, 19394, 1939b). 
The work of others were reviewed by Masloy 


: 5 A aad ial Survey, South Australian Depariment of Mines, Rox 38, Rundle Street UO, Adelaide 


Trans, R. Soc. 5, Aust, Vol, 96, Part 2, 31] May 1972, 


68 W, OV. PREISS 


(1960), Mast of these uuthors tacitly accepted 
the validity of a formal binomial nomencta- 
ture Por stromatulites. 


Cloud (1942) was the first to question the 
validity of such uw classification, arguing that 
stromatolites are built by associations of alyal 
species. Similarly, Johnson (1966) has more 
recently rejected the use of this nomenclature, 
and suggested rather that only actual algul spe- 
cits should be named. if they are present. 
Nevertheless, Rezak (1957) found it useful to 
retain a binomial nomenclature and used the 
defined tuxa successfully for intrabasinul corre- 
lution. 


Since the controversy regarding the classi- 
fieation of stromutolites arose, ut Jeast three 
schools al thought have evolved, Firstly, a 
small group of Russian students (c.g. Vologdin 
1962), like Johnson, considered that. oniv 
actual ulgul remains can be validly named, But 
algae are very rarely preserved in Precambian 
stramutolites, and mast of the micro-structures 
referred hy Vologdin to fossil algae are very 
doubthully of organic origin. 


A second proup rejects the concept oF bio- 
logical comirol over stromutotite morphology, 
and uses purely descriptive classifications to 
aid environmental interpretations. For example, 
Masloy (1960) used “generic” names such as 
Collenia, Conpphyton and Glebrilella, but he 
modified these by a series of descriptive Latin 
auljectives. Logan, Rezak & Ginsburg (1964) 
used symbols, and formulae to deseribe various 
features of stromatolites, which they showed to 
be influenced by the local environment. Both 
Maslov's multinomial nomenclature and the 
variable descriptive formulae of Logan ef al, 
tend to be cumbersome, and cannot in them- 
selves describe all the useful characters of 
stromatolites. Some of the simpler formulac 
are. hewever. very useful in routine lichd des- 
criplions, Hofmann (1969a) found difficully in 
upplying a binomial nomenclature to the surno- 
matolites of the Gunflint Iron Formation, and 
later (Hofmann 1969b) discussed the signifi- 
eance of various characters of stromutolites, 
concluding that dilferent morphologies are 
moze likely to be environmentally than bio- 
legically controlled, Tloffman (1967, 1969) 
gave an outstanding example of the use of 
stromatolites in palaeocurrent determination 


The third school is a Russian group which 
describes and classifies stramatalites on tbe 
Basis of morphology and microstructure. and 
uses them for blostratigraphy, Their first results 


were reported by Keller et al, (1960). Despite 
differences Of emphasis today between dilferent 
workers, all usc a binomial nomenclature with 
the form taxu “group” (analogous ta genus) 
und ‘form™ (analogous to species), They have 
found that the time ranges of the defined taxa 
are restricled, and this allowed them to sub- 
divide and correlate Late Precambrian sections 
throughout much of the USSR. The biosirati- 
graphy was supported by numerous radiometric 
datings, both K-Ar determinations on glaueo- 
nites and K—Ar. Rb-Se and U-Th-Pb deter- 
minattons on intrustves. The subdivision is ws 
follows: 


Cambrian 
$70 = LO my. 
Vendian 
———— —- 6380 = 20 my 
Late Riphean 
-—- = 950 = 50.m,¥, 


Middle Riphean 

1,350 = 50 my. 
Early Riphean 

1,600 = 50 mv. 

The approach of this group was applied to 
Austruliun strontarolites, and it was found that 
many of Ihe Russian taxu do occur here, in a 
similar order of succession (Glaessner, Preiss 
& Walter 146% Preiss 1971). The resulting 
correlations with the daicd Russian sequences 
were in agreement with most of the radiometric 
evidence available for the Australian Precam- 
brian. 


The successful use of stromatolites in bto- 
striligruphy implies that their morphotogy a al 
least partly controlled by genctic characters of 
algue which evolve in time. The concept of 
biological control is supported by some studies 
of madern algal mats (Eardley 193%; Hom- 
meril & Rioult 1965; Monty 1967). Each of 
these suthors. has shown the partial dependence 
of mat type on the predominating algal spe- 
cies present. This in turn affects the miicrostrue- 
ture and lamina shupe of the stromatolite, und 
indirectly, the pross morphology. Thus, decid- 
ing which characters are genetically determined 
and which are directly shaped by locul environ- 
mental fuciors becomes the major difficulty in 
classifying stromatolites. 

The Russian work of recent years has showa 
that it is mainly the columnar stramatolites 
which are of value in biosirutigraphy, Only 
Komat (1966) has given 4 detailed account of 
laterally linked stromatolites but their useful- 
ness has not been confirmed io the extent of 
thar of columnar forms. In this study, atten- 


SOUTH AUSTRALIAN STROMATOLITES 6y 


Fig, DIAGNOSTIC TERMINOLOGY 
MODE OF OCCURRENCE 


BIOHERMS: SUS, Cow Aa Kean SN 
= (a — _————————— on ES 
SUBSPHER ICAL DOMED TABULAR 


NNN rs ete Pips 
2 mS 


PEAAANIMEO GLAD Dg eter IL 
Fonte Oe TE > 
OORT A ere all) Y wy 
Ree TT ARE TTT 

S S a anual 


BOMED 


EUAN DURUCUTE Lt a URAC CAR ASLAN UA cb cd AAA 
COTTA OTT 


BIOSTROMES: TABULAR 
BRANCHING AND COALESCING 


nay Gel 


alpha beta gammo SLIGHTLY 
PARALLEL DIVERGENT MARKEDLY DIVERGENT COLUMNS 


COLUMN SHAPE AND MARGIN STRUCTURE 


— 
NICHE AND 
PROJECTIONS PROJECTION RIBS WALL 
m 
LAMINA SHAPE pn 
GENTLY CONVEX RECTANGULAR “WAVY /\ EX 
STEEPLY CONVEX ‘RHOMBIC —- WRINKLED UNCONFORMITY esta ESA 
CONOPHYTONS 


NON-COLUMNAR STROMATOLITES 
SS RES an 
SS ERBReESe 


ee aa fs ge oN Ne 
oo OO Pe 
FLAT-LAM INATED ASS SRS SE Zeer 
ae oN el NNN 
= = = SOS SSS 
OAS ENS Undulatary Pseudocelumnor oS 
CUMULATE ; LATERALLY -LINKED COLUMNAR-LAYERED 


Fig. 1. Diagnostic terminology found useful in the description of stromatolites, The diagrams illus- 
trate features discussed in the Appendix. 


70 W, V. PREISS 


Hon was also concentrated on columnar forms 
since these have the most characters allowing 
them to be classified, Therefore the binomial 
nomenclature has been applied onty to these. 


The terms used here to describe stromptolite 
characters, ate largely based on translations of 
Russian lerms, with minor alterations and addi- 
wens. Mast of the new terms introduced by 
Ilofmann (1969h) are unnecessary from the 
point of view of this study, The diagnosti¢ ter- 
minuloxy proposed by Glagssner er al, (1969, 
Fig. |) bas been expanded (Fig, J), and the 
terms used in the descriptions ure defined in 
the glossary, Appendix I, 


‘Taxonomy 


Ia general the methods of stromatolile study 
and classification used by the Russians. Krylov 
(1963, 1967), Semikhatov (1962), Nuzhnoy 
(1967) und Komar (1966) have been applied 
here, inclucling hinomial nomenclature. Al- 
though muny of the group names have been 
accepted! tn pulaeohotanical literature os 
penera, of. Baicelid, Conophiyron and Gyiino- 
yolen by Andrews (1970), it is considered that 
retention of the terms greup and ferm empha- 
sizes the distinction between stromatolites asx 
organo-scdimentary structures and actual [rag- 
mentary plant remains to which the terms 
form genus and ferm species may be applic- 
able. While similar groups of characters are 
studied Cor cuch stronvitolite, The relative 1ako- 
none significance attached to any particular 
Character muy vury from luxon to ison, tle- 
pending on its diagnostic value, Mostly. groups 
are defined an the basis of gross morphology. 
column shape, branching and margin structure, 
Lamina shape and microstructure are fre- 
quently useful in the distinction of fornrs But 
sometimes these features are diagnostic at 
group level—Conopiyion, lor example, is diag- 
nosed by its lamina shape, and is characterized 
by purecuhirly distinet Januinulion, Baicalia 
tends Lo he characterized by banded laminating, 
except where altered by cliagenesis, Although 
the presence of a wall is a dingnostic difference 
between sone eroups, Walter (1970) 7 has des- 
etihed single stromatetiles which are unwalled 
in their flower parts and walled at the top. in 
which case other features are diagnostic. No 
similar situation is known from South Austra- 
lias, 

IL is considered unnecessary to use categories 
higher than the group, as Raaben (1964, 


1969a, 1969b) has done. Her higher taxa are 
somewhat arbitrary and several alternative ones 
could be proposed, but all are equally question- 
able. On the other hand, the wodery asa sub- 
division of the fornt is Gseful in cases where 
finer subdivisions can be made, and is there- 
fore retained for Coaaphyton garganicnm. 


It could be argued that a single name would 
be sufficient ta characterize a particular stroma- 
tolite, but the value of a binomial nomen- 
elature is that it mudicates real similarities and 
differences between various forms. Thus groups 
contain one or more forms which all share a 
number of characters considered diagnostic for 
that group. Forms are distinguished within a 
vroup whenever there are suflictent gross or 
microstructural differences, But the essential 
compunson between closely related Turns 
would be lost without a binomial nomenclature. 


The chief difficulty in the Waonomy of sino 
matolites is the isolation of discrete character 
combinations, where intergradution is commean- 
Thus, subjective choice may he required in 
some causes. Where the morphology temsains 
uniform Vhreughoul a particular occurrence, & 
single name can easily be applied, but if there 
is Varialion within the occurrence, the delini- 
ton must be broadened, Whether or not the 
morphology of 9 stromatoltte [rom another 
egcurrence falls within this range of variation 
is dificult wy decide. Coaversely, if a signili- 
cantly dillerent morphology occurs as a disercte 
portion of un deecurrence, is this to be classi- 
fied separately? Examples of stromatoliies with 
a broad cunge of variation ure Puagassta etina 
and Linelln muanyallina, Toth of these show a 
spectrum of intergrading branching types and 
colunin shapes, even within single outcrops, se 
that the range of variation between specimens 
of different areus lies within the range of varia 
on in one locality, and these are therefore 
included in the one form. 


Tr has been found that many characters over- 
lap. and distinctions must be made even al 
group level Gn the most Commonly occuring 
expression, ie. the mode. of each character. 
This is especially true of branching, Bexania is 
characterized by «-paralie] and some §-parallel 
branching, while ysparallel is rare. In Gymno- 
sole +-parallel predominates, but not to the 
total exclusion of the other types, Similarly, 
there is overlap between the branching styles 
of Burealia {the forms of which show a Cromen- 


Unpublished PAD. thesis, University ef Adelaide. 


SOLTH AUSTRALIAN STROMATOLITES 7 


dous vanation of branching, as shown by 
Krylov 1967) and that of Tungussia, Bul while 
Baicalia has predominantly slighthy to mode- 
rately divergent branching, in Twngessia mae- 
kedly divergent branching predominates. 
Although it is often easy to recoynize groups 
on the basis of even limited reconstructions and 
lonvitudinal sections, the identification of forms 
is more difficult and subjective, Forms are dis- 
tinguished orn minor features of column mor 
phology, lamina shape, or microstructure. 
Microstructure is the most difficult chataciter 
la use, partly because different types intergrade 
to some extent and partly because it ia so easily 
altered by diagenesis. The distinctive lamina- 
tion. of Conopiryron is somewhat ¢xccptional, 
and is amenable to statistical analysis. Although 
Raaben (1969s) has attempted similar studies 
on Migeria and Semikhatovy, Komar & Serebrya- 
‘kov (1970) have measured the sizes of clots 
and pelleis in Bexonia, it is uncertain Whether 
or not the structures measured are primary. 


The laminae of most South Australian 
columnar branching stromatolites are too 
diffuse and variable to allow a detailed statis- 
tical study, although the well-preserved repre- 
sentatives of the banded microstructure of 
Baicalia burra might be amenable, 


Stromatolites can be classified only on the 
basis of combinations of characters, and, as 
Walter (1970, unpublished) has also con- 
cluded, the classificatory significance of charac- 
ters must vary to some extent from taxon to 
taxon, The classification has been found em- 
pirically to be useful in that the resulting taxa 
are lemporally restricted. The questian arises 
as to the fundamental meaning of these taxa, 
and why they wre sa restricted, Several possi- 
bilities. exist: 

(1) Each form is built by a particular asseo- 
ciation ef algal species, and forms 
change as the content of the associations 
changes. 


Each form is built by a dominant algal 
Species, in association with other species 
that have Jittle effect on stramatolite 
morphology. 


(3) The environment. and not the algal 
composition, entircly controls the stro- 
matolite morphology. 


ff (3) were true, we should expect 4 tem- 
poral restriction of forms only if the environ- 
Ment has systematically evolved in time. Lt is 
difficult to sce how local factors such as current 
activity or sediment accumulation which could 


(2) 


conceivably control stromatoliie morphology, 
can exhibit continent-wide, if not world-wide, 
Unidirectional change. On these grounds, this 
possibility must. at present be rejected. If (i) 
were true, Wwe could expect the morpholazy te 
change gradually as the overall algal coniposi- 
tion changes. one species replacing another in 
the association. On the other band, if one 
specics controls the morphology, a rapid change 
would be expected. At present, itis not possible 
to fell which of (1) and (2) is correct and 
possibly both upply,; the first possibility may 
explain the intergradations sometimes observed 
between taxa, When classificahoa becomes difii- 
cult. 


Although Hofmann (1969h) regarded Cone- 
pPhyren as apatt from other stromatolites, 
recent work has shown that conuphytons pos- 
sibly intergrade With columnar branching 
forms such os Baicalia (Shapovalova L968). 
Similarly Bertrand (1968) described intergra- 
dations of Conephyren and branched forms. 
While the taxonomic significance of these 
changing morphologies and theit relationship 
to environmental factors has not. been fully 
determined, it is clear that Conophyton is not 
fundamentally different from other stromato- 
lites, 

It is concluded that stromatolites must be 
defined on combinations of characters, the sige 
nificance of each of which may vary in differ- 
ent taxa. The fact that some taxa have much 
broader ranges of variation than others results 
from the necessity of grouping intergradina 
morphologies present in single stromatohte 
occurrences, 


Methods 


Stromatolites were studied both in the field 
and in the laboratory, but ficld observations 
were often limited by outcrop conditions and 
lichen cover on tock sutfaces. Where possible, 
the mode of cecurrence, column shape and 
umangement and branching were observed in 
order to vjin an impression of the tutal yari- 
ability, 

The variable nature of stromatolites neces- 
silates sampling of sufficient material to dcter- 
mine the modal expressions of churacters pre- 
sent. Depending on the size of columns, large 
specimens weighing from 4-70 kg were col- 
lected, and the relative position and orientation 
were noted, Ideally, bioherm centres and mar- 
gins were both sampled. 


The diagnostic gross features of colummar 
stromatolites can only be determined from a 


72 W. V. PREISS 


three-dimensional view of the stnicture, This 
is achieved by the method of “graphical recon. 
struction” described by Kryfov (1963), A 
series of LO to 15 serial Jongitudinal slabs 2 to 
6 mm wide were cut on an oil-cooled 60 cm 
diamond saw with a saw cut about 2 mm wide. 
‘The columms were outlined jn pencil on the 
slabs and traced on to a block diagram frame- 
work on tracing paper. each Jongitudinal sec- 
tion heing parullel to the front. face of the 
block. The reconstructions were retraced with 
shading to show surface morphology and finally 
redrafted by stippling. 


Lamina shape, margin structure and micro- 
structure were studied in large, longitudinal, 
thm sections, up to 20 em long, Thew thick- 
ness varies with the nature of the rock, hut in 
general they must be thicker chan petrological 
sections to preserve the distinctness of the 
structures. Carbonates were mostly identified 
by staining with Alizarin Red §, but were [re 
quently checked by X-ray diffraction powder 
photographs, 


Systematics 


For each group from which forms are des- 
cribed, 4 diagnosis, a list of the known cansti- 
tuent forms and the stratigraphic and geogra- 
phic distributions are presented. Fotms are 
diagnosed only if described here for the first 
ume. Descriptions are given under the headings 
made ef occurrence, colunu shape and arrange- 
ment, branching, margin structure, laenine 
shape and microstructure, The interspace sedi- 
ments and the nature of secondary alteration 
aré wiso described since they provide importunt 
clues to the depositional environment and dia- 
genelic history, 


The distribution of forms refers to both their 
geographic distributions and te the rack-strati- 
graphic units (Thomson er «il. 1964) in which 
they occur. Reference ts made for each locality 
and stratigraphic unit to the relevant geological 
sheet (either 1:63.3A0 er 1:250,000 map 
sheets, Geological Atlas of South Australia). 


Tt was found convenient (Preiss 1971) to 
subdivide the Adelaidean into two time units: 
the Barly Adclaidean, represented by all seci- 
ments up fo the pre-tillite unconformity, and 
the Late Acelaidean, represented by sediments 
from the base of the lower tillite to the base 
of the Cambrian. This subdivision reflects both 
@ climatic change and a major change in stro- 
matolité assemblages. Ages of stromatolites 
will be referred to as Early or Late Adelaidean, 


but the probable corrélations with the subdivi- 

signs oF the Riphean will he noted in each case. 

Type specimens are kept in the Department 

of Gealogy and Mineralogy, University of Adec- 

vide, catalogued under numbers prefixed by S, 
Group ACACIELLA Walter 


Walter has supplied the group name Avca- 
ciella and the following diagnosis, 
“Type Form: Acaciella australica (Cryptozeen 
dusivalicum Howchin 1914). 
Diagnosis; Nearly straight, parallel or radially 
arranged sub-cylindrical columns with a-, f- 
and rately y-parallel and very slightly divergent 
multiple branching. On column margins are 
numerous low bumps and occasional small cor- 
nices ond peaks; small areas Of wall occur in- 
frequently, Laminae dominantly are rect- 
angular. rhombic of gently domed and are not 
markedly wavy or wrinkled; the microstructure 
is streaky.” 
Content: Acaciella australica Walter, A 
angepere f. nov. aid A, aneusta f. nov. 
Age and Distribution: Adelaidean to Early 
Cambrian: Loves Creek Member of the Bit- 
ter Springs Formation, Central Aust.; the 
Lower Cambrian of S. Aust.) the Wundowje 
Limestone and Brighton Limestone equiva- 
lent. Umberatana Group. S. Aust. and as 
erralics in the lower (Sturtiqn) elaciily, 8. 
Aust, 


Acaviella angepena f, nov. 
FIGS. 2 10, lla 

Material Forty-seven specimens from ie 

localities. 

Nolorype; S460 (Pigs. 2a, 10c), Lower 

Cambrian, | km south of Angepena MS., 

Northern Flinders Ranges, 

Name After the type locality, 
Diagnesis: Acaciella with vertical or radially 
arranged columns. or pseudocolumus, which 
may branch upwards from either flat-laminated 
or small cumulate stromatolites.. Columns may 
branch upwards into minute, irregular columns, 
Bridging 1s. extremely common. Microstructare 
is regularly banded, with thin continuous 
Jaminae. Verniform microstructure may be de- 
veloped, 


Description 
Mode of oecurrence: Cambrian stromatolites 
were studied in outcrop only in the Angepena 
aren, where lenticular stromatolite beds consist 
of closely spaced ellipsoidal and domed bio- 
herms 3 to 50 m wide. These overlie flaggy, 


SOUTH AUSTRALIAN STRGMATOLITES am] 


laminated, dark grey limestones with irregular 
efosional contacts. Cumulate or  pseude- 
columnar stromalolile individuals commence 
growth upon the erosional highs, aod pays up 
inte radially arranged or parallel short columns 
with very numerous bridges, At bioherm mar- 
gins, columns and psetidyceltimns become hori- 
vontal. and laminae are deflcxed parallel to the 
overhanging sides of the bioherm. su thut here 
growth actually proceeded duwnwards (Fig. 
1Ou & b, showing longitudinal sections of a 
bioherm margin), Where adjacent biaherms 
become contiguous, they are overlain by a 
damed biostromal layer of columnar, pseudo- 
columnur und columnar-layered stromatolites, 
similur to those of the bigherm. Ac the edge of 
a stromatolite bed, the terminal hioherm has an 
abrupt vettical margin and the laminae bend 
downwards only slightly. The surrounding sedi- 
ment of dark Jime mud accumulated synchran- 
ously with stromatolite growth, and vecasional 
lgul Jaminae are intercalated with it: the bia- 
hero probably never hod more than 10 em of 
relicf over the surrounding sediment surface. 
Also, there is evidence of contemporanedus 
compaction of the time mud the bicherm resis 
upon (Fig, 10ce). the lower layers of the sur- 
rounding muds ate depressed, while the upper 
ones simply abut agalnst and cover the bio- 
herm, 


Column Shape anid Arrangement: Column 
shape is highly variable in single bioherms, 
mainly due to different degrees of coulescing 
and brilging. The structures vary from laterally 
linked pseudocolumns with some discrete small 
cumuli (Fig. 10d) to frequently bridged and 
coalescing calumas (Fig 11a), to discrete, 
parallel subcylindrical columns (Pigs. 2d, f, a: 
10f). The latter chiefly make up Muawson’s 
(1925) callection from Italowie Ciorge. In all 
specimens where columns are reasonably dis- 
erete, they are smooth to slightly bumpy, some- 
limes wilh pointed terminations (Fig. 2b, j), 
while others branch into minute columns 1 to 
Jom wide. Columns are commonly Jess than 
! em diam., but brogd, cumulate columns up to 
10 cm diam. have heen observed. Transverse 
sections of columns are round. reunded poly- 
sonal or lobale (Fig. 2d, f. 2). Columns may he 
verticul or radially attunged, especially on dhe 
margins of contiguous bioherms. Dolomitiza- 
tion of interspaces frequently obscures the ori- 
ginal margins of the minute columns sq that 
their shape cannot be accurately devermined_ 


Branching; Branching is most commonly g- of 
B-parallel, vccasionally y-parallel, Columns fre- 


quently branch into narrower columns which 
do not regain the former diameter. Some 
branches ure in the form of thin pointed pro- 
jections (Fig. 2j) Ac bioherm margins, 
branching may remain parallel] (Fig, 10¢) or 
hecome radial (Fig. 100), but here the stronia- 
tolites are Jargely pseudocolumnan 


Margin Srracture; Colum matgins are rarely 
preserved intact, Commonly they are corroded 
by dolomite thombs, if the interspaces are dolo- 
milized; otherwise very fine stylolites may be 
developed. Bridging is extremely common in all 
specimens except those from Matvsan's cal- 
Iection from Ttalowie Gorge. in which the 
columns are mostly disctete. These alsa have 
the smoothest margins, with only slight, acca- 
sional bumps and ribs. Columns are always 
unwalled. the laminae thinning only slightly 
near the columo murgin, Laminse may slightly 
overhang the margin, but long peaks and ¢or- 
nices are absent, 


Lonine Shape: Fig. 8a illustrates common 
lamina shapes: most ire gently convex. Of 101 
Jamina measured, 69% have height to diameter 
ratios (b/d) beneath 0.2 and 0.4, only 7% 
have ratios greater than 0.6 (Fig, Say, Laminue 
ure smoothly domed, without sharp changes in 
shape from lamina to Jaminoa, A few of 
Mawson's specimens from Itulowie have wavy 
laminae, Of Wavelength 3 tc 10 mm, amplitude 
1 10 3 mm (Fig, 10f). 


Microstructure; Microstructure in all spect 
mens is regularly, thinly banded, with contin. 
ous Jaminac of uniform thickness across & 
colunin width. In most specimens there is little 
contrast between dark and light laminae, except 
in the amount of organic pigment. Some speci- 
mens, especially fram Angepena, have ifregu- 
larly tubular, sinuous, agastamosing. vermi- 
form spurry patches, O05 to 0.1 mimi thick anc 
up to 0.6 mm long, crossing the dark Juminae. 
Dark laminae, varving in thickness from 0,03 
to 0.07 mm, consist of xenotopic calcite of 
grain size varying from 0.003 to 0.01 ming, 
stained With vroy orgunic pigment, bul in some 
specimens, subhedral dolomite rhombs of grain 
size O=.1) to 0.02 ram are interspersed, Minor 
subangular quartz, silt may be present, Indivi- 
dual laminae ure continuous, anc af constanr 
thickness across the colunin width, but may be 
markedly wavy, In specimens with veentiform 
microstructure, dark laminae are generally 
thicker. up te 0.3 mm, but remnunts of finer 
luminution ut ullen preserved, The hwundaries 
of the sparry patches are often irregular and 
their orientation varies from petpendichlsr to 


TA W. V. PREISS 


gently inclined to the lamination, but is com- 
monly at a high anule to it. The vermiform 
microstructure may be consistently developed 
preferentially on one side of u column Trans- 
verse sections of the tubules ate found to elon- 
gated, irregularly Oriented and anastomoxing. 
The tubules may be interpreted as algal borings 
in the fine, lime mud laminae, but not the 
whole sediment was alfected. since homo- 
geneous ahd bored! laminae occur side by side 
This fact also makes it unlikely that they arc 
casts of actual algal filaments. Whe distribution 
of borings on one side of columns may be 
environmentally determined. Bathurst (1466, 
p. 20) illustrated a sequence of events Invalved 
in boring by algac; if the process were stopped 
ul stage 12), and the borings infilled’ with 
sparry calcite, a structure similac tw the vermi- 
forn) microstructure olf A. angepena would 
result, Light laminae are 0.03 to 0.1 mim thick, 
frequently indistinct, but continuous ucross a 
colimm width. They are especially poorly dif- 
feruntiated in specimens with vermiform micro- 
structure, Where the tubules may pass ocross 
the light-dark lamina boundaries. Light laminae 
consast of xenolopic calcite, often with inter- 
locking ervstals G.015 to 0.03 mm in diameter. 
Subhedral to) cuhedral UO. mm dolomite 
rhombs are scattered throughout the light 
linninac in some specimens, 


Jrevspacey> Tnterspaces are filled either with 
Altered micrite or fine sandy and silly micrite 
Specimens from Jralowie (Mawson's collee- 
tion) have very narrow interspaces Filled with 
sparse, qngular quartz silt. supported by a 
Micrite matrix (Fig. 1Of), somelimes exten. 
sively dolumitized, with inequigranular hypidio- 
topic dolomite ringing in gruin size from O.005 
10 O.L mm. Extremely finely disseminated hac- 
mutité may be present tn interspaces. Strom.- 
lolites from Angepena also have sandy inter- 
spaces, byt these are more frequently inter- 
rupted by bridging laminae, Subangular to sub 
rounded quartz grains vary in diameter from 
0.08 to .S tom, and may be partially or wholly 
replaced hy calcite, Qpids and small intraclasts 
occur very rarely. 


Secondary Alteration: Dolomitization is com- 
mon jn all specimens, and is probably of late 
diagenetic origin. Within columns, rhombs post- 
dule the vermiform microstructure and have 
also formed in the micrite of interspaces. anc 
in places, interspaces may be totally dolo- 
mitized, Here sparry calcite occurs as irregular 
patches between dolomite rhombs perhaps fill- 
ing @ secondary porosity, The grain size and 


density of dolomite rhombs decrease murkedly 
across the ealumn margins: perhaps interspaces 
were uriginally more porous, lo cause the pre- 
ferential dolomitization (in Fig, MWe¢ nore the 
dark calcite columns and the white, dolomine 
interspaces), StvJolites may follow column niat- 
gins. or may be grossly cross-cutting. Haema- 
tiie dispersed through carbonate a4 probably 
sccondaty. In fealowie specimens, it is concen- 
trited in interspuces which pass inte ine stylo- 
lites. Minute irregular cal¢ite veins cut the 
whole rock, apparently predating the major 
dolomirzation. Large patches of course sparry 
cnleite wire bounded by markedly lobate fine 
atvlolites, suggesting their origin as solution 
cavities, 


Compariseny 


In sross morphology (mode of occurrence, 
column shape, branching and margin structure) 
the strumatutites from Ttalowie are similar to 
AcecieHa Waller, Columns are less discrete in 
uther aureus, due to frequent bridging and 
coulescing, but their columnar portrons ure 
similar to those of Ttalowie specimens. Micro- 
structures are uniform, except fur the local ver- 
mniform structure interpreted as ulgal bering, 
Madiyonites nuiwsont Walter, from the Middle 
Cambrian Jay Creck Limestone of the Amua- 
deus Basin, also has vermiformy microstrucnure, 
but here the tubules ure more consisiently deve- 
loped, and wre complesty intertwined. the inter- 
yening micritic areas being reduced to clots. 
The gross form of Muarlivanites anivsonl is 
similar to same Acuciella aaxepena in having 
numerous ireegular frequently bridged columns 
und pseudocoluniis, however, it lacks the sub- 
eylindrical, parallel branching, discrete columns 
Yourd at Tralowre. Acacrelle — ueyeprentiea 
resembles Verella svehhasiea Krylov in having 
evenly banded lamination and wide columns 
branching into nurrow columns, bul has ragged 
eclumo margins and Jacks the wall of P- 
uethbayitd, Mlicta composite Sidoroy is siniilue 
jn also possessing vermifarm microstructure, 
hut is distinguished by its very smooth, walled. 
ehlurins. At thes stage uf is difficult to be cer- 
tun of the content of the form Aeuetella ange- 
pena, Despite some variation nf column shipe 
(specimens frony [talowie have predominantly 
suhevlindricail, discrete columns, while those 
from Angepensa have numerous bridges and less 
recular column margins, all the specimens 
studied are included in the one form, since 
these column morphologies intergrade and the 
Microstructures rempin constint. The stremp- 
tolites are assigned to the group Acetyl on 


dratielle angepena, from Lower Carshrian Timestancs, !'lliiders Ranges, 14) —Molptype, 5460, 
| km $. of Angepena H.S,) (b)—S458. 1 kin) S. Of Angepena ALS.y (c). (2) & (i }—S489. 1 
kin S, of Angepena WS; (0) & tz) SB. 4.8 km W, of Itatowie Gorge (oallecred hy Sir 
Douglas Mawson}; (f)—S44, 4.8 kin W, of Nalowie Gorge (collected by Sit Douglas 
Mawsan?; (h)—Probable 4. angepena, near Old Wirrealpa (collected by Mr, P, G, Haslett): 
(i)—Possihle 4. dagepena, 4.8 km WL af Tiulnwie Gorge. 


V6 WwW. ¥. 
the basis of gross morphology, They are dif- 
ferentivied Trom other forms of the group by 
their thin, continuously banded microstructures 
and by frequent. development of bridges and 
pseudocolumns. The very narrow, minute 
columns into which broader columns branch 
are abseac in other forms 
A ferrupinous specimen from Old Wirrealpa 
is problemutical. Its dark laminae are strongly 
haematiti¢, the hacmatite being in part distri- 
buted inte minute dendrites. The small columns 
branch from basal cumuli, the interspaces being 
filled with recrystallized biomicrite fhyolithius, 
spange spicules. archacocyathan and brachio- 
pod fragments may be recognized). Although 
the grass morphology resembles that of Aca- 
cielld ungeperia (Piz, 2h), the extremely regu- 
lar lannnation is atypical of stromatolites, and 
the possibility of an inorganic orgin for the 
structure cannot be excluded 
Divrifucion: Widespread in the dark lime- 
stones of the Lower Cambrian at Angepena, 
Old Wirtealpa, near Point Well, at Mern 
Merns, Beliana Hill, Chace Range, near 
Narina HS,, Moro Springs. south of Balca- 
noona, and 4.6 km west of Ttalowie Gorge; 
Flinders Ranges, South Australia. (COPLEY 
and PARACHILNA 1:250,000 map sheet 
areas. } 
Age: Early Cambrian. 


Acaciella angusta £. noy. 
FIGS. 3a-m, 11d-f, 12 

Material; Thirteen specimens. trom two loca- 
lities plus eight spécimeéns: of Uncertalyy iden- 
tificulion From a further two localities. 
Ffoloiype, S401 (Figs 3c. e 120), Brighton 
Limestone equivalent, Depot Creek, South- 
crn Flinders Ranges. 


Name: After the city of Port Augusta, 32 
km south of the type occurrence. 


Diagnesiy: Acuciella with extremely frequent 
coalescing and bridging of columns at all levels, 
and with broad und narrow columns closely 
associated, Colutin margins bear short ribs, 
low bumps and short cornices. Laminae are 
geutly to moderately steeply convex or rec- 
angluar, and of distinct, revulerly streaky 
Microstructure. 
Description 

Made of OQecurrencée: The stromatohtes form 
lenticular and tonguing bicherms (Fig, 12a) 
varying in thickness from 3 m fo 50m, and cx- 
tending laterally for up to nearly 2 km, inter- 
calatéed at varying siraligraphic levels within 


PREISS 


the Brighton Limestone equivalent, Most com- 
monly. growth commences on a substrate of 
ooid wod intruclust grainstones, as laterally 
linked stromatolites, up to 3 m thick; these gra- 
dually develop interspaces to form broad, 
bridging and coulescing columns (Fig. 11d). 
At various levels, these columns branch into 
harrawer columns 1 to 3 cm wide, frequently 
with parallel hasal and slightly divergent upper 
branches (Fig. 12c). Occasionally, narrow 
columns anse directly from an undulatory or 
fat-laminated base (Fig. 11f). Columns 
repeatedly alternate with continuous undula- 
tory Or flat-laminated stromatolires, which com- 
monly intertongue with the adjacent sediment; 
they apparently mark periods of reduced influx 
of coarse sediment. At bioherm margins, 
columns bevome slightly inclined. Rarely. there 
are hemispherical bioherms with columns 
strongly inclined at their margins, 

Column Shape and Arrangement: Basal 
columns are up to 20 cm wide, of ipegular 
shupe, with frequent coalescitig and bridging, 
Their margins are frequently inclined. although 
laminae remain subhorizontal. The narrow 
columns are 1 ta 3 cm wide, and up to 10 cm 
long between branches (Figs. 3a-j; | 2b-d). 
Transverse sections are round, rounded poly- 
agonal, clongated, or complexty lobate, At least 
some of the clonggtion is of tectonte origin 
Columns are Straight of gently curved, with 
slight swellings and constrictions (Fig, 4a-j); 
a few are short and marrow, and terminate their 
growth after a few centimetres (Fig. je). 
Coalescing is an Tréquent thut almost all 
columns are interconnected! one specimen con- 
tains numerous irfegular, short, frequently 
bridged and coalescing columns. 


Branching: Branching is frequent at all levels, 
and generally multiple (Fig. 3a-j). Broad basal 
columns divide by a-parallel branching into 
narrower columns, which frequently branch 
ugain at intervals of less than 10 em} this 
branching is usually q- or 8-parallel, occa- 
sionally y-parallel, or slightly divergent (Fig. 
3e-c, g). Near points of coalescing, branching 
tends to be more irregular; gamma-purallel or 
divergently branched columns approach each 
other and coalesce (Fig. 3d). 


Margin Structure: The \ateral surfaces of all 
columns bear relatively low bumps, short dis- 
continuols ribs, and a few peaks and comices 
(Fig. 3a-j). In places. bridges, varying from 
delicate bridges only onc or two laminae thick 
to massive, thick bridges (Fiz. 3g), are very 
frequent; in other places, columns remaln rela- 


SOUTH AUSTRALIAN STROMATOLITES 77 


uvely unilfected by bridging throughout most 
of their jength. Columns are unwalled, und be- 
tween bridges and cornices their margins are 
relatively smooth. Depending on the degree of 
convexity. Jaminac approach the margin at 
various angles. 


Lamina Shape: Lamina shape varies according 
to columin diameter; narrow columns have 
moderately convex, Or sometimes steeply con- 
vex, laminae. h/d greater than 0.6 being rare. 
Broad columns have very gently convex to rect 
anvular laminae (Fig. 8b). Of all laminac mea- 
sured, 70% have h/d between 0,2 and 0.6 
(Fig, 9b), Laminae are most frequently 
smooth, but sometimes brojdly wavy, ¢spe- 
cially before branching. Laminse frequently 
become donbly-crested before branching (Fig. 
\2b-d), but the interspace so formed may be 
heidged over, in Which case the colunin resumes 
its former growth pattern. 


Mirrasiructure: I the best preserved speci- 
mens, distinct, regular light and dark (gzeen) 
laminae, and in places macrolaminae up to 4 
mm tnck, alternate, forming a regular streaky 
microsiructure (Fig. 12). Durk larmincde, vary- 
ing In thickness from 005 mm to 2 mm, are 
smooth to gently wavy, occasionally wrinkled, 
and have parallel ipper and Jower boundaries. 
Single laminae have relatively constant thick- 
ness across the column width, but frequently 
lens out They consist chicfly of hypidiotopic 
to idiotopic dolomite, of grain size ranging 
from 0,005 to 0.02 mm. The crystals are equi- 
dimensional, commonly euhedral, and stained 
pale green, which gives the laminae. their 
colour. Dolomite crystals are densely packed in 
the dark luminae. leaving only occasional irre- 
gular undolomiuzed patches. vonsisting of 
xenotopic calcite, ranging if grain size from 
0.003 to 0.01 mm. Light lamina: vary in thick- 
ness [rom (07 te 2 mm, single laminae having 
constunt thickness. They are sparsely deolo- 
mitized, and consist of xenotopic ta hypidio- 
tipic calcite, varying from 0.01 to 1.02 mm in 
grain size, with scattered euhedral dolomite 
rhombs, 0.005 to 0.04 mm long. Laminae are 
frequently grouped into broad mucralaminae, 
up to 4 mm thick, in which very thin, lenti- 
cular’, erther light or durk laminae predominate. 
In places, laminae are slightly wrinkled, or 
uraped over underlying irregularities; in one 
case laminae are domed over Jenses af spatry 
calcite, probably open space fillings (Fig. 11e). 
In a few places small scour structures up to 
2 mm decp are cut into the tops of dark 
laminae. Occasional euhedral to subhediral red- 


dish hrown limonite peains of 1.01 to 0,02 mm 
diameter (possible pseudomorphs after pyrite) 
eceur in both lamina tWpes, 


Qiterspaces: The distance between columns 
vanes from | to 14) mm. Interspaces are filled 
with banded limestone, layers of micrite 1 to 
6mm thick alternating with thicker intervals of 
partially dolomitized intramicrite, Laminag im 
the interspace commonly abut against the 
column margins, having accumulated after the 
arowth of that part of the column (Fig, 12c) 
The micrite laminac, consisting of xenotopic 
calcite of grain size varying Sram 0,003 to 0.01 
mm. are frequently silty, and slightly graded, 
Senerslly with sharp upper boundaries, and are 
overlain by matrix-supported intramicrites and 
some aomicrites, This sediment may originally 
have been more porous, as It is extensively 
dolarmitized; the dolomite is of similar texture 
Lo that in columns, All remnant calcite is re- 
crystallized to a hypidiotopic sparry masaic; no 
micrite matrix remains. Alternatively, this cal- 
cite may fepresent infilling of volds left by 
Solomitization, Intraclasts, which may bea pre- 
served as undolomitized micrite, or entirely 
idiatopic dolomite, are from 1 co 10 mm long, 
and up ta | mm thick, and may represent 
eroded fragments of algal mat- Strongly reerys- 
tallized dolomitized oolites are actasionally 
present. Tnotraclasts. which communtiy [te at a 
high angle to the bedding, muy have fine 
erained laminae draped over them. Coarse sedi- 
ment mux was periodic; columns may have 
had up to 2 cm of relief over the interspace 
sediment or a bridge, then the interspaces were 
filled rapidly with intraclasts and finer ¢al- 
ca/cous ‘sediment. During periods of |elative 
quiescence, lime mud accumulrted to form thin 
layers. In some specimens, bridging is very fre- 
quent, so thal there never was more than ahnut 
a centimetre of relief. 


Secondary Alrereasian; Liile is preserved of the 
primary difference between the light and the 
dark (green) laminge, which now differ in the 
extent of dolomitization, The dolomite is equi- 
granular, idiotopic, and probably secondary, 
although a detrital origin cannot he miles? ot 
fi the dolomite originated by replacement of 
calcite, the preferential dolomitization of dark 
laminae may indicate that they were oristhally 
more porous, Smull irregular patches of 
coursely crystalline sparry calcite within both 
columns and interspaces post-date dolomitiza- 
ton, and are associated with fine calcite veilis. 
Stvlolites aze very fare, being restricted 16 a few 
which afe concordant with ihe liminaiion or 


74 


column margins. The grec staining of dolo- 
mite crystals oxidizes under subaserial weather- 
ing to form finely disseminated limonite, which 
may be concentrated along colunin margins or 
slylolites. Colamas are commonly slightly fiat- 
tened parallel to an axial plane cleavage, which 
is better developed south of Depot Creek and 
at Munialilo Creek. ‘The cleavage is an irre- 
gular fracture which passes around, not 
through, slromatalite columns and is commoaniy 
expressed as stylolites in the carbunate rucks. 
A specimen from Mundallio Creek contains 
light laminae wiih prominent radiating, struc- 
tures; these consist of dolomite crystals aligned 
in Tows almest perpendicular to the Juaminution 
(Fig. 12d), and may represent a dolomitized, 
earlier ueicular texture. In specimens from the 
Wundowie Limestone (Wundowice Bore and 
Copley), column margins have been ulmast 
completely removed by stylolites, Icading to 
unceruinty of identification (Fig. 3k. 1 & m). 


Comparisons 
The predominantly parallel branching (o- 
parallel at hase, then 8- or rarely y-parallel) 
and almost total absence of a wall, identify the 
stromatulites as Acuerelle. 


4eagiella qugusta is distinguished fron A. 
oumniica by the rarity of discrete, brogu, basal 
culumos. by its mode of oecurrence (lenticu- 
Jar and wonguing bioherms instead of tabular 
tnd domed hiostromes), by its extremely fre- 
yuent coalescing and bridging, and by its very 
distinct mierostructure, The mode of ogcur- 
renee and microstructure abe distinguish it 
from A dnewpena, A. angusta has many wavy, 
sometimes Tenticalar laminae, and prominent 
macroluminac, the dark laminas being pre 
ferentially dolomitized, A, augusta is very simi- 
lar in gross morphulugy to Eucapsiphora para- 
disa Clond & Semikhatoy, from the Puradise 
Creek Formation neu Mt, Isa, N,W, Queens- 
and. &. paradisa is difficult to distinguish on 
the basis of the published description, but 
apparently has a patchy wall. 

Specimens of poorly preserved stramutolites 
from the Wondeowie Limestone near Copley 
and Wundowie Bore. originally tentatively 
identified as Linella munyallina (Preiss unpubl.) 
are belter assigned to Acacielfa augusta on the 
basis of column shape, branching and micro- 
structure. Where column murgins are not re- 
moved by stylolites. they arc unwalled, 

Diswibuyion; Brighton Limestone, Depot 

Creek ard Mundallio Creek. Southern Flin- 

ders Ranges, and possibly the Wanclirwie 

limestone near Copley and Wundowie Bore, 


wlsy.: 


PREISS 


Northem Flinders Ranges (PORT 
AUGUSUA und COPLEY 1:250.000 map 
sheet areas), 


Age: Late Adelaidean, correlated with the 

Late Riphein of the USSR, 
Acaciella. form indet. 

FIGS. 3n-y; 11b, ¢) 
Material: Two specimens from one locality. 
Description 

Mele of Occurrence: Both stromatolite speci- 
mens are erratic boulders in the lower 
(Sturtian) glacials; their provenance is un- 
known. 
Column Shape and Arrangemens One speci- 
men (S509). eunsisty of pseadocolumns ancl 
frequently bridged columns, oriented sub- 
parallel to shebtly rachialing, wn passing tuter- 
ally inte flat-laminated stromatolites (Fig. | le). 
The uther specimen (8339) consists of rather 
smooth, erect, parallel. cylindrical, discrete 
columns, I-5 cin wide. Transverse sections are 
round ar rounded polygonal {Frg. 3n-p). 
Brunching: Hraneching ix commonly a- or p- 
parallel; columns either retain their width or 
widen gradually before branching. Axes of 
branching columns mav be very slightly diver- 
pent (Fig. 11b), Specimen §539 shows only 
dichatomous branching, but S509 has sume 
multiple, o-parallel branching. 
Murgin Stractvre; S539 hos wo ruther smtoth 
margin structure. with low bumps and a few 
very shorr peaks and overhanging laminie 
(Fig, 3n-p). There is no wall; laminae simply 
tefminute, withoul appreciable thinnmg, at the 
column margins. Bridges are extremely fre- 
quent in SSOQY, but otherwise column: margins 
are similar to 3539, Few columns in $509 are 
entirely discrete. 
Tamina Shape; All laminae are gently conven 
(Fig. 8c). hid never exceeding 0.5. und 84% 
of Jaminue measured have h/al between, 0.2 and 
4 (Fig, Yel. Laminae are smoothly curved, 
rarely rectingular, and without wrinkles or 
sharp Nexurcs. Occasionally Inminae are slightly: 
wavy, and belere branching always develop 
multipte crests (Fig. 11b, ¢), Laminae are not 
normally deflexed at rhe column margins. 
Microstructure: Microstructure consists of very 
smooth or broadly wavy, light und dark. dolo- 
mitic, striated ta banded, laminae. ‘There is little 
conttust between laminac. Dark laminae are 
0,05 jo 0.5 mm thick, and commonly pinch 
and swell slightly across the column, and in 
places they ate lenticular, but otherwise, they 
have smooth, parallel boundaries. They consist 


(a)-(); Acactella augusta, Brighton Limestone equivalent, Umberatana Group, Flinders 
Kanges: fa), (bo, (d¢) & (1) S404, Depot Creek, Southern Minders Ranecs; (cl & fey— 
Holotype, 5401, Depot Creek: (£) S538, Mundallio Creek, Southern Flinders Ranges; (2) & 
(h)—S396, Depot Creek: (j)—8537, Depot Creek: (Kk), (1) & (m)—Paossible 4, augusta, 
Woadowie Limestane Member, Woudowie Bore. Northern Flinders Ranges; (n). fo) & (p} 
—S539, Aenelella F, indet. from an erratic in the Sturtian glacials, N.E. of the Pnorams 
Diapit, (q)—S509, Acaciella F. indet,, from the same locality. Sketch traced from a thin sec 


tion, 


80 W. V. PREISS 


ot pale vrey stained hypidiotopic dolomite of 
gruin size varying from 6.003 to 0.015 mim. 
Light laminae vaty in thickness from 0.05 to 
1.0 mm, generally with little change across a 
colummn. thinning only shghtly towards colunia 
margins. They. consist of hypidictapic to idio- 
topic transparent, unstained dolomite of grain 
size varying from 0.015 to 0.06 mm. Very 
characteristic of $539 is the presence of very 
fine, limonite-rich solution surfaces, concordant 
with the laminac, Although these are probably 
stylolites, surfaces with lithe or no wrinkling, 
which follow the fine-scale structure of laminae 
exactly, are especiully common (Fig, 11b). In 
places these are only about 0.5 mm apart, and 
light Jaminae muy be separated by them, with- 
out intervening durk laminae. 

Interspaces. Interspace sediment 1s completely 
dolomitized, consisiing of equigranular, idio- 
topic dolomile of grain size ranging from 0.01 
to 0.05 mm. There is little contrast between 
columns und interspaces, but small amounts of 
subangular quartz silt ure present in the inter- 
spaces, Fraginents of slightly darker stained 
dolomite, of similar texture to the matrix, pro- 
bably represent original intraclusts. The nature 
of the matrix cannot be determined, bul the 
sparsity of intraclasts stiggests that these were 
mud supported, Intraclusts are better preserved 
in S509, 


Secondary Alteration: Dulomitization af the 
siromatolites and interspaces is clearly second- 
ary, as indicated by the general idiotopic,, equi- 
granular texture, and poor preservation of the 
finest structures, Stylolites are of at least two 
generutions: the earliest stylolites are almost 
perfectly concerdani, without lobes or wrinkles: 
these possibly predate the dolomite euhedra, 
which in places cut inte them, and certainly 
predate 2 relatively coarse grained dolomite 
vein (grain size up to 0.1 mm). The vein is 
itse]E cut by more prooounced, slightly diseord- 
ant stylolites. Occasionaly cross-cutting srylo- 
hites cut interspaces and columns, some follow- 
ing column margins. Dotomitization almost 
certainly predates the erosion und depasition of 
the clasts into the glacial sediments. 


Comparisons 

The straight, chielly 4- to f-parallel branch- 
ing unwalled columns allow assignment to the 
group Acaciella, They are clearly distinguished 
from Acueella uvgasia by the discrete, rather 
smooth, more cylindricul columns; although 
bridging and coalescing oceur in S509, this 
specimen is considered to represent the basal 
purt of the stromatolite bed. The distinct, sub- 


evlindrical columns with telatively smooth mar- 
gins and gently convex laminae are similar to 
A. awsiralica Walter, but the specimens are 
inadequate for identification, 

Distribution, As clasts in the lower 
(Sturtian) glacials, on the Aynks of Enorama 
Diapir, 6.4 krn North of Orgparinna HS., Cep- 
tral «ss Flinders Ranges {PARACHILNA 
£:250,000 map sheet area). 

Age: Probably Adelaidean, but net younger 
than the Sturtian glacials 

Group BAICALIA Krylov 

Bailie Krylov 1943+ 64. Semikhatoy 1962: 

198. Komar 1966: 82. Krylov 1967: 25. 

Nuzhnev 1967: 135, Cloud & Semikhatay 

T1969: 1035. 

Type Form: Baicalia baiealica (Muslov) 

Krylov, from the Cluntuy Suite of the Pri- 

baikalye [based on Collenia baicalice Maslov 

19374, 287]. 

Diagnosix: Tuberous, bumpy, swelling and von- 
stricting. parallel to markedly divergent branch- 
ing columns, genetally withour wall, with fre- 
quent overhanging laminae. Lamination is dis- 
tinctly banded. 

Coment; B, baicalica (Maslov) Krylov, &, 
Kirgiste Krylov, B. rata Semikhatav. &. 
unea Semikhatov, B, prima Semikhatov. B. 
ampla Semikhatov, A, meilensis Nuzhnov, 
8. maica Nuzhnov, 8B. ainica Nuzhnov, B- 
minnie Komar, 8. eapricornia Walter and 
B. burra fF. nov. 


Age: Middle Riphean to carly Late Riphean. 


Baicelia barra f. nov. 

FIGS. 4, 5, 6, 13, 14, LSa-c 
fietcalit spp. Glacssner, Preiss & Walter 
1969-1056. 

Marerial: Thirtythree specimens [rom ten 
localities. 
Holotype: S222 (Figs. 3a & d), Skillogalee 
Dolomite 3.2 km west of Yatina, Southern 
Flinders Ranges, 
Nanie: From the Burra Group in which the 
stromatolites occur. 
Diaginus: Baicalia vith moderately frequent 
slightly to markedly divergent branching, irte- 
gular, coalescing columns with highly variabJe 
lamina shape and continuous, distinctly bandetl 
microstructure, 


Description 


Mode of Occurrence: Two modes of ocvur- 
tences have been noted: biostromal and bhio- 
hermal. the latter occurring only at one locality 


SOUIH AUSTRALIAN STROMATOLITES $] 


(Yatina), Hiustromes vary in thickness Jrom 
0.3 ta 2m, the stromutolites being eventy dis- 
tributed throwehoul their extent; they have been 
followed for |00 mm or more, without lensing, 
before the outcrop disappears under soil cover. 
Biostromes are frequently interbedded in green 
shales (e.g. Myrtle Springs, Willouran Ranges), 
platy dulomiles (e.g, Arkaroola, Worumba) or 
massive dolomites (e.g. Burra). The bicherms 
at Yatina afe restricted to (wo thin beds; they 
are small lenticulac stromatolitic mounds, 
approximately 20 to 30 em thick and up ta 
1m wide (Fig 130), interbedded with and 
surrounded laterally by platy and shaly dark 
grey dolomites, The overlying sediment is 
draped over the mounds, showing that the 
stromalolites had at least 10 cm relief over the 
surrounding surface. Colunins arise from sub- 
strates in several ways: (1) Flat-laminatesd 
slromufelite passes gradually up inte undula- 
tory and pscudocoluimnac stromatolites, then 
Info discrete, vertical to inclined columna, often 
with steeply domed lieminac fe.g, Burr, West 
Mount Hut); (2) Colurnns urise directly fram 
eroded surfaces of laminated or intractastic 
dolumites (e.g, Yatina, Fig. 44))3 (3) Columns 
arise from flat-laminated stromatolite wine broad 
cumuli (e.g. West Mount Hut), The degrees of 
discreteness of columns varies greatly; in some 
beck. columns are almost immediately bridged 
over by Jaterally linked stromatolites, but 
usually columns remain discrete for 20 lo 30 
cm. In some arcas new sets of columns may 
arise from pscudocolumns. The upper surfaces 
of biosuwomes vury From flat (¢,g, in the Wil- 
louran Ranges, Burra) to broadly undulating 
(e.g. Worumba). 


Column Shape end Arrangement: Columns are 
tubctous, varving From subeylindrical ta irregu- 
lar, with round, oval and irregular cross sec 
tions (Figs. 4, 5, 6), Elongated or Aattened 
colunms are varionsly onented. The diamerer 
of colarns varies Erom J to 10 ¢m, most com- 
monly 3-5 em, with rapid swellings and con- 
striclions, Coltinins aré 2-15 cm high between 
branches, Some but not all columns are con- 
stnicted at the point of branching (Figs. 4c, f; 
Sc, d). The onientation of columns varies 
greauly From vertical to inclined, and is some- 
times sub-horizontal for short distances (Figs, 
4e, 5)). Column axes vary from. straight to 
strongly curved. In some specimens, the upper- 
most columns swell matkedly upwards und be- 
come bridged over by laterally linked stroma- 
tolites. Adjoining columas coalesce very fre- 
quently, even in the discrete portions, but speci- 


mens from Burra show the least coalescing and 
bridging. In the Willogran Ranges, column 
arowth is frequently mterrupted by penecon- 
[éemporaneous erosion; columns may grow over 
hroken-off fragments of eurlier columns, con- 
tribuung to the irregularity of the structure. 


Branching: The most common form of hranch- 
ing is moderately divergent (Fig. 4a, i, 6F) 
though same sub-parallel branching (Fig, 4e, 
fg, m) and some very markedly divergent 
branching occtirs (Figs. 4a, Sd & 7)- In same 
specimens several] branches arise from nearly 
one point (Fig. 4a). Branching is moderately 
frequent, the length of column between 
branches commonly being only a few centi- 
metres; bed at any one point of branching it 
is usually dichotomous of less offen trichoto- 
mous, In some specimens branches arise at a 
high angle to the main columns, and then tumn 
shurply upwards, Some columns arise from the 
side of a main column (Fig. 14d). Great varia- 
tion is seen even in single outcrops, 


Murgin Srreciure: The lateral surface varies 
fromm smooth to very irregular, laminae ap- 
proaching the column margins at Yaridus 
angles. Some specimens have very patchy 
walls, while the intervening unwalled areas are 
smooth or only slightly fringed with smull 
peaks and cornices, for example those from 
Burra (Fig. 14d), Yatina (Fig. 4a}, River 
Broughton (Fig. Sc), Arkaroola (Fig. Sd, ¢). 
Willouran Ranges specimens contin both 
smooth and highly irregular edges, with large 
overhanging peaks composed of one of niore 
laminae (Fig, 6b & c}_ Frequently large swell- 
joys are composed of numerous Jaminae over- 
hanging a constricted portion of g column 
(Fig. 4¢), Bridges between columns are espe- 
cially common ncar the tops and bolioms of 
biostromes (Fig, te), 


Lamina Shape: The lamina shape is most com- 
monly genily convex, but varies in single speci- 
mens from very gently convex to pearly cons 
cil; nutny laminae jire sleeply convex, Micro- 
unconformities are espectally prominent in 
specimens from the Willouriun Ranges, bul 
occur ta some extent in alf areas. In places, 
branching cammences upon a partly eroiled 
column surface (Fig, 15b). Fig. $d illustrates 
the more commenly occurring lamina shapes; 
92% of lamina have h/d between 0.1 and 0.6, 
the mode being h/d between 0.3 and 04 
(28% 1 (Fig. 9d). Generally, the widest 
columns have the most gently convex laminae, 
while strongly elongated columns hive laminue 
gently convex in the sectinn parallel to the long 


eo W. V. PREISS 


Fig.4 


Fig. 4. Baiculia burre. Skillogalee Dolomite. Burra Group, Southern, Flinders Ranges: (a) & (c)— 
Holotype. S222, 3.2 km W, of Vatinn: (b)—S218. same locality; (d) SIS{, 13 km S.W, of 
Werumba H.S. (e) & (f) S151. same Jocalityy ¢g), ( & (()—S221, Dutton’s Trourh H.S,, 
i¢ km S. of Burm; (h) & (1) —S314, same locality; (kJ—S534. sume Jocality; (o1)—float 
specimen, River Broughton, W, of Spalding. 


as. 0 


Baicalia burra, Skilogalce Dolomite, Burra Group: ().—S333, Dutton’s Trough H.S., 14 km 
S, of Burra; (4)—S534, same locality, (cj S383, River Broughton, W. of Spalding; (d)— 
§456, 6.4 km S, of Arkaroola; (¢)—S457, same locality: (f)—S491, 2-4 km E. of Myrtle 
Springs H-S. (upper member of Skillogslee Dolomite); (g)—S489, same locality; (h)— 
$490, same locality: ((}—S488, 1.4 km E. of Myrile Springs HS. (lower member of Skillo- 
ealce Dolomite); (j)—S487. same locality; (k)—S319, the Avoodale Mine, Lyndhurst (col- 
lected by Mr, P. J. Binks); (1)—S302, West Mount Hot, Willouran Ranges; {m)—S99, same 
locality (collected by Mr. C, R. Dalgarno): (m)—S97. near Chintapanna Well, Willouran 
Ranges (collected by Mr. C. R. Dalgarno). 


Pig, #- 


Baicalla burra, Skillogalee Dolomite, Burra Group, north-western part of the Adelside Geo- 
synctine: (a), (b) & (f)—S96, Chintapanna Well, Willouran Ranges (collected by Mr. C. R. 


Dalgarno}; (c)—S98, West Mount Hut, Willouran Ranges (collected by Mr C. R. Dulgarna); 
(t)—S496, 4.8 km W, of Copley; (2)—S301_ West Mount Hut, Willouran Ranges, 


SOUTH AUSTRALIAN STROMATOLITES Ha 


axis. nd steeply convex ut right angles to it, 
Rarely co laminue turn over sharply and thin 
al the column margins, to form a wall. Geoe- 
rally, where a patchy wall Js present, ii is 
formed by the edges of steeply convex or paru- 
holic laminae (Fig. 14c), Frequently, laminae 
develop two e¢rests, anticipating branching 
immediately above (Fig. 4a, m). On a smaller 
scale, lamina shupe varies from smooth and 
regularly curved to slightly wavy, with discon- 
tinuous curvature and sharp crests. Both types 
occur in single specimens (Fig. 14). 


Microstructure: The microstructures and tex- 
tures Ohserved tn the clifferent areas vary con- 
sidvrably depending oo the deyree of recrystal- 
lization. In the best preserved specimens, the 
layering comprises alternating relatively thick, 
contmuous, very distinct. light and dark 
laminae, giving a banded appearance. Some 
are single homogeneous thick layers, while 
others are macrolaminac consisting of several 
very thin light-dark Jamination pairs, Most 
commonly single laminae traverse the whole 
eolunia width, excepr where cul by micra- 
unconformities. Onids or other detrital grains 
may be included in the laminae, Upper and 
lower boundaries of laminae are usually 
smooth and even, sumetimes wavy or broadly 
wrinkled, bul always more or jess parallel. 
Fxeéeptians occur only where erosional scour 
has taken place during growth, Rarely, lenti- 
colar swelhngs occur. Aight Javinae vary in 
thickness from .02 ro OS mm, very rarely to 
1.0 mo Mast light laminae thin towards the 
column edges, but rarely Jens out. In the best 
preserved specimens, the sparry dolomite forn- 
ing them is ineqnigranular, xenotopic, and of 
vrain size ranging from ().005 to 0,06 mm. 
With greater recrystallizatlon an equigranular 
mosaic of 0,05 to 0.2 mm grain size results 
fee. Burra), The light laminae usually have 
sharp and smooth upper boundaries, hut some- 
limes grade down into grumous textured lami- 
hac, consisting of irregular and interconnected 
mivritic patches vp to 0.1 mm diam,, set in 
xenolopice =equidimensional sparry dolomite 
wilh a grain size of about 0.01 to 0.03 mim, ic. 
partially recrystallized’ dark laminac. In some 
specimens (2.g. Yatina, West Mount Hut, 
Worumba}, the light [aminae contain detriral 
granules. including small fiat inteaclysts, up to 
0.5 mm tong, and rare onids up to 0.3 mm in 
diameter, Overlying laminae are draped over 
the larger detrital grains, Laminae in the Cop- 
ley specimen may be pelletal (Fig. !4e}. Dart 
faminae occur either singly, alternating with 


light Jaminae, or in dark macrolaminae, Thin 
dack laminae are commonly 0.04 lo 0.3 mm 
thick, but macrolaminae range up ta 2.5 mm 
in thickness, generally constant across the 
column, or thinning slightly towards the mar- 
gins. They are either continuous, or consist of 
a-series of aligned lenses, each up to 0.2 mm 
long. In well preserved specimens the dark 
laminae have smooth, sharp, parallel boun- 
daries; rarely, single laminae may be wrinkled, 
suggesting intrafarmational crumpling during 
growth. Well preserved dark luminac consist of 
dense, brownish-pigmented xenotapic dolomite, 
of equidimensional grains 0.003 to 0.0L mm 
dism,, but vertical and lateral gradations from 
unaltered to erumous textures are commen. 
Where dark laminae are grouped into miacto- 
laminae, they alternute with very thin, discon- 
tinuous light laminae, and {requently fuse to 
form solid. thick dark Taminac. 


Interspaces, A tow specimens have interspaces 
filled predominantly with bedded dolomite mnd 
(e.g. Burra), but generally the sediment is 
unhedded intraspucite or oosparite, less com- 
monly intramicrite, Frequently, intraclasts are 
derived from the erosion of  stromatolitic 
columns; in places, 9 large fragment torn from 
a column has acted us a base for new growth. 
Tntraclasts ate flat to gently curved tabular 
dolomite pebbles up to 3 cm long, | lo 2 mm 
thick, and only slightly rounded. Many are 
Fragile and could have survived very litle 
transpor!, They contiin the typical internal 
laminations of the associated stromatolites, and 
are probably derived directly from them. Gcca- 
sional flat pebbles stand vertically, bur ene- 
rally they lie fiat or imbricated, Ooids yary in 
shape from round to oval, 0.2-[.0 mm diam, 
und consist of one dark-rimmed sparry Liver 
coating a micritic core, or Jess commonly. 
sevetal sparry layers. Most commonly, allo- 
chems are closely packed and cetnened by a 
clear, sparry dolomite cement, Some specimens 
contain significant amounts of dolomite mud, 
variously recrystallized. forming a macrex be- 
tween allochema; in these cases the seciment is 
poorly laminated. 


Secondary Alteration; Secandury alteration has 
extensively modified the textures and often the 
microstrictures af stromatolites from many 
drews. The following four stages of alteration 
miy be fecognized: 

(1) Peneconemporaneous. The fact that dolo- 
mite consistently constitutes the whole rock to 
the exclusion of calcite, while still preserving 
fine structures, suggests very early dolomitiza- 


86 W. V. PREISS 


tion, during the growth of the stromatolites, It 
is also possible that penecontemporaneously 
Uolomitized lime muds were reworked and 
trapped in the algal mats. During growth, cro- 
sion by strong currents scoured the living sur- 
faces of columns, creating micro-unconformi- 
ties. In some specimens (e.g, West Mount 
Hut), laminae may be separated by lenticular 
vughs, later filled with sparry dolomite (Fig. 
12c), These voids were probably formed by 
arching up of laminae, perhaps due to lateral 
expansion in growth of the algal mats building 
the siromatolites, of 1 partial desiccation. 


(2) Early Diaveneric. Black chert very com- 
monly replaces porlions of slromatolites and 
interspace sediments, Sometimes dark laminae 
are preferentially silicified, perhups during 
growth, but more commonly, silicification post- 
dites the growth of the colamns (e.g, one side 
of a column may be replaced). In places, silt- 
cified laminae are brokea by minute dolomite 
filled cracks, 


(3) Late Disyvenetic. Dark Jaminae and macro- 
laminae may be recrystallized tay grumous tex- 
lures, Consisting of patches of dark, dense 
micritic dolomite (remnants. of the original 
cutbonute) varying greatly in size from 0.005 
to 0.1 mm, set in a matrix of xenotupic sparry 
dolomite, of cquidimensional grains ranging in 
size from 0.01 to 0.03 mm. Light laminae are 
commonty slightly recrystallized and sparry, 
consisoug uf hypidiotopic to idiotopic. equi- 
dimensionul dulomite grains of similar size to 
(hove of the sparry matrix of the grumous tex- 
lures, Coarsely recrystallized laminae also 
occur, i, places Cutting ucross. the fine stmuec- 
ture of primary laminar and corroiling their 
boundaries, They consist of idiotopic tran- 
sparent. dalomele of grain size up to 0.1 nun. 


(4) Yeerwnte. The only specimens affected by 
tectonic deformation are from the Burra 
region, Here columns are slightly Nuttened and 
laminae ate erenulated along a slight tectonic 
foliation, These are alsa the must highly meta- 
morphuscadl, cisplaying the greatest degree of 
recrvstallization, Tensional joints filled with 
toursely Crystalline Uolumile are commnan it 
most areas. 


Conpartrons 
The stramatolites are assigned to the group 
Buicalia on the basis of their taberous, swell- 
ing and constricting, humpy, variously oriented 
columns general absence of wall, numerous 
overhanging peaks and short cornices, and 
gencrally divergent branching. Some specimens 


have horizontal colomns for short distances 
fesombling Fusigussia, but are distinguished by 
the absence of the multiple horizontal branch- 
ing characteristic of Tunpussia, and hy their 
generally oainre mgged column margins. 
Baicetia busra is distinguished from B. printa 
Semikhatov, 8. aimica Nuzhnov, and 8. cupri- 
cornia’ Walter, by fits Frequently divergent 
branching and general complexity of columns, 
and from &, mimeéa Kormar by its larger size 
and more complex structure, Some specimens 
resemble &. baicalir (Mastov) Krylov, but 
most have more inclined and irregular columns. 
8. lacera, B. rare, B. ample ans B. wit Semi- 
khatoy are not adequately ijlustrated for reli- 
yble comparison, and the illustrated micro- 
structures are badly altered; single specimens 
of B. burre may show microstructures similar 
to B, wnea, B. lacera, and especially the pelletal 
laminae of B. rare, Some specimens have long 
overhanging peaks and thus resemble A, 
ingilenvis Nuzhnov, but are distingunshed by 
more frequent and divergent branching. &. 
hurra most closely resembles B. rare Semi- 
Khatey and 8, maica Nuzhooy) it is distin- 
suished from &. rara in that neither pellectal 
tuminse nor knee-shaped bends in columns ure 
consistently developed, and from B. meied by 
its more inregwlar and coalescing columns, and 
its more continuous kiminae, 
Disteibution: Widespread in the Skillogalee 
Nolomite, Burra Group: Yutton’s Vrough 
H.S., 16 km south of Burra; Scrubby Raoge, 
27 km south of Burra; 3 km west of Yatinat 
River Broughton, 8 km west of Spatding: 11 
km south-west of Worumba; 11 kn south 
of Arkaroola; 3 km west of Copley; 3 km 
east of Myrtle Spnngs HLS. near Leigh 
Creck: West Mount Hut, 27 km west of 
Witchelina H.S. wod Chintapanna Well, abouc 
If kny west of Witchelina DLS. Possible 8. 
burr oceyrs also in the Skillogalee Dolo- 
nite, Depot Creek, but chesé have not been 
studied in detail. Specimens from possible 
River Wakefield Group, Carnieton (Fig. 15c) 
ate inadequate for identification, but are 
possibly to he inchided (BURRA, ORRO- 
ROO, PARACHILN’. COPLEY. ANDA- 
MOOKA and CURDIMURKA 1:250.000 
map sheet areas), 
Age. Early Adetaidean, correlated with the 
younges| Middle Riphean of the USSR- 
Group BOXONIA Korolyuk 
Boxonia Korolyuk 1960-129 Komar 19Af: 
7¥. Cloud & Semikhatov 1969:1036- Glaess- 
ner, Proiss & Walter 1969: 1056 


SOUTH AUSTRALIAN STROMATOLITES 87 


Type Ferm: Boxenia grorilis Koarolyuk. 

from the Bokson Suite, Eastern Sayan. 
Diagnosis; Straight, subevlindrical columns 
with moderately frequent g- to @-patullel 
branching and smooth, walled margin strac- 
ture, 

Content; B. gracilis Keorolyuk, 8B, lissa 
Komar, B. krasivica Golovanoy. B. allab- 
jueice Komar & Semikhatoyv, B. ingilice 
Komar & Semikhatov, B, hiunca Raaben 
and B. perruknarre Walter. Raaben 
{1969a) places B. erumulosa Komar into 
partial synonymy with & gracilis Koro. 
lyuk, &, diverata Sidorov has only 2 patchy 
wall and may therefore be exchided- 


The South Australian form is Bexenia 
melroxa. 


Age; Late Riphean and Vendian. 


Boxonia melrosa f nav, 
FIGS. Fah. |5d-f 

Meaverial: Four specimens from one locality. 

Nelotype: S503 [| Figs. 7b, e & dy 15e), 16 

km west of Melrose township, Southern 

Flinders Ranges, 

Nunes After the type locality. 
Diagnesix: Baxania with long, narrow, ¢lasely 
spaced columns, a- and A-parallel braiching, 
without very broad basal columns, with occa- 
sional rounded projections, and with indis- 
tinctly banded. muderitely convex, Ipminae 
lacking pelletal microstructure. 


Description 

Mode of Occurrence: ‘The stromatolites are 
relatively poorly exposed in a faulted area, so 
that relationships are not clear, At least two 
bioherms occur, preserved as grey or pale buff 
dolomite. ‘I'he beds are overturned, dipping 
south at about 40°, The narrow, parallel 
columns arise difectly from futetally linked 
stromatolites. partly pseudocolumnar, the base 
of which is not exposed. The overlying 
columnar portion ts approximately 6 m thick 
and constats of vertical columns neal the centre 
of the bioherm, and inclined colunms at the 
mareios, where they pass laterally into pseude- 
columnar stromatoliies. Columns are overlain 
hy wavy laminated stromatolites, which cover 
the whole bioherm, Bioherms are of cumplate 
shape, broadly domed, up to 60 m long, and 
are surrounded by flut-bedded dolomite, 


Column Shape and Arrangement: Columns are 


straight, erect, subcylindrical. smooth to gently 
bumpy, with circular or slightly lobale, 


rounded polygonal cross-sections, 1-5 cm diam. 
(Pig. 7a-h). The diameter of 2 single column 
generally remains constant throughout its 
length. Columns may reach a length of up to 
20 cm between branches, bul some columns 
are only a few centimetres high, occasionally 
in the form of rounded projections 

Branehing: Branching varies from a- to f- 
parallel; y-purallel branching is tate (Fig. 7h). 
Commonly a 3-5 cm column divides into two 
or three narrower, parallel, very closely spaced 
columns, I-2 em diam, (Fig. 7e,d,f), Ooca- 
sionally, two narrow columns may coalesce 
{Fiv, 7c). Not all branches develop into long 
columns; some terminate their growth only a 
few centimetres above branching (Fig, 7d), 
Mirgin Structure: The lateral surface is even, 
smooth ar with low, broad bumps, ap to 
several centimetres wide. with a relief of I—5 
mim. Peaks anc cornices are entirely absenr, bur 
very rarely bridges up to 1 cm thick occur 
between adjacent columns. A multisaminate 
wall is almost ubiquitous, At the margins of 
columns Jaminae are poorly preserved, bur in 
plices up to 10 laminae may be seen to com- 
prise the wall. Single laminae gencrally extend 
for au distance of 1-2 em down the column 
margin (Figs. |Sd—f). 

Camina Shape; Laminae are most commonty 
moderately convex, hemispherical, in places 
approaching rectangular (Pig. 8e). Frequently 
they are slightly asymmetrical, especially in in- 
clined columns, Belore branching, lammae 
usually develop two crests, The degree of con- 
vexity, h/d, is moderately conslunt, even in 
columns of differing widths, Of laminac mea- 
sured, 91% have h/d between 0.3 und 0.7, the 
mode (39%) being 0,5-0.6 (Fig, 9¢). The 
shape of crests varies from tightly urenate to 
gently rounded (Iig. Se). Most laniinae are 
broadly wavy (wavelength up to 8 mm, ampli- 
tude 1-2 mm) but not wrinkled, 
Microsinneture: Microstructure is poorly pre- 
served in both pale and dark specimens; 
laminac are broadly continuous, with smooth, 
patullel upper and lower boundaries, but niwy 
he broken into a series of clots and lenses by 
recrystallization, and even where their con- 
tinuiry is preserved, they are extensively em- 
bayed by recrystallized carbonate. Microstrac- 
ture is indistinctly banded with alernating 
darker and lighter laminac. Light jane vary 
in thickness from 0.08-0.4 mm, but usually 
thin towards column margins. Continuity és 
usually retained across a column, although the 
finest Jaminac: {frequently Jose their identity by 


88 


recrystallization, The laminae consist of tran- 
sparent, slightly incquigranular (of grain size 
0.01-0,04 mm) cguidimensional dolomite of 
polygonal, hypidotopic fexture. Within this 
occur irregular 0.05-0.1 mm segregations of 
darker, greyish pigment, with no relation to 
grain boundaries. These ate apparently reni- 
nants of pigment left by partial recrystallization, 


W. ¥, PREISS 


as they may grade into more or Jess continuous 
laminae, Distinct round to oval pellets (as in 
Russian Boxonia) are ubsent. Dark laminae are 
less continuous, and often diffuse. Their thick- 
ness vaties from 0,08-0.3 mm; towards the 
margins they trequently thin or lens out com- 
pletely, and do not take part in the formution 
of the wall. (The Jayering in the wall is 


Fig.7 


Fig: 7. 


Koxonia melrosa, Brighton Limestone equivalent, Umberatuna Group, 1.6 km W, of Melrose: 


(a), Ce), (2) & (h)—S502; (b), (ce) & (d)—Holotype, $503; (1)—S504. 


% 


a} 


? 


Dy») 


fm 


Representative examples of lamina shape: (4)—Acuciella angepena: (b)—A. augusta: (cj— 
Acaciella f. indet.. (d)—Haicalia barra, (€)—Boxonia melrasa. 


SOUTH AUSTRALIAN STROMATOLITES ay 


extremely indistinct). In places, they lens out 
also within the central patt of a column, Dark 
laminse are composed of équidimensional, 
Renotopic, equigranular dolomite (of grain size 
ranging from 0,003—0.01 mm), and in places, 
ure disrupted into a series of irregular clots and 
lenses separated by sparry dolomite. 


Interspaces: Interspaces between columns are 
extremely narrow (usually Icss than 5 mnt), 
und are filled with partially recrystallized dola- 
mite mud, now largely of fincly grumous tex- 
ture, containing in places, round or ovoid clas- 
uc pellets, 0.2-0.7 mm diam. Much of the sedi- 
ment ts vaguely Jumingted, the laminae abutting 
against the walls of columns, which they post- 
date. 


Secondary Alteration: Stromatolite columns 
and interspaces consist of dolomite, considered 
to result from the replacement of original cal- 
cium carbonate, Most fine structure has been 
lost; dark laminae are outlined mainly by seg- 
regations of dark pigmented dolomite, but re- 
crystallization has partly embayed and partly 
obliterated the fine dark laminae. The irregular 
distribution, of pigment is due to recrystalliza- 
tion. In places, coarser, sparry laminae of grain 
size up to 0.08 mm occur, and may contain 
dismembered remnants of dark laminae. Stylo- 
lites are moderately frequent, and usually dis- 
cordant to the Jamination. In places they follow 


Fig.9 


column margins for short. distances, removing 
the wall. Occasional thin dolomite veins follow 
the path of stylofites. Some stylolites are 
parallel to overall bedding, and displace column 
axes slightly (Fig. 7c). 
Comparisons 

The stromatolites are assigned to the group 
Boxonia on the basis of their long, smooth 
walled columns with moderately frequent a- 
and B-parallel brunching. Katavia Krylov and 
Acaciella Walter have similar gross structure; 
Katavia is distinguished by its very prominent 
humps, while Acaciella generally lacks a wall. 
Minjaria Krylov also has parallel straight 
columns but is distinguished by its less frequent 
branching. Most other described forms of 
Boxonia have well defined pelletal microstruc- 
tures; forms are largely distingutshed on the 
basis of the size of the pellets, A specimen of 
B, gracilis sent by M. A. Semikhatoy and I. N, 
Krylov, has pellets consisting of rounded car- 
bonate grains with dark, fine-grained rims. 
These are absent in B. mielrosa, which also has 
less wrinkled laminae. 8. melrosa is distin- 
guished from B&. ingilica Komar & Semikhatov 
by its ubiquitous wall and straight columns: B. 
allahjunica Komar & Semikhatov apparently 
has some complex branching, B, lisse Komar, 
B. gracilis Korolyuk, B. grumulosa Komar, B. 
bianca Raaben and B. krasivicu Golovanoy 


Histograms of Jamina convexities. The convexity of a lamina is ihe ratio of the height of 
that lamina to its diameter (h/d). Histograms are plotted for each stromatolite form at inter- 
vals of 0.1; # is the number of measurements made for each form: (a)—Acaciella angepena: 
(b)—A. augusta; (c)—Acaciella £. indet.: (d)—Baicalia burra; (e)—Boxonia metrose. 


9D W. V. PREISS 


may all be synonymous. & melrose is distin- 
guished from 2B. pertaknurra Walter (in press), 
which also. lacks a pelletal microstructure, by 
its more steeply convex laminae, its occasional 
short, projection-like columns and by the 
absence of well defined broad basal columns. 
B. melraxa most resembles B. fissa, from which 
it is distinguished by the absence of pellctal 


microstructure, and by the presence of some 
short, projection-like columns. 


Distribution: Brighton Limestone equivalent, 
1.6 km west of Melrose (ORROROO 
1:250,000 map sheet areu). 


Age: Late Adelaidcan, correlated with the 
late Riphean of the USSR, 


Acknowledgements 


Tam indebted to Prof. M. F. Glavssner for 
supervising this study al the University of Ade- 
Inide, and to Dr. B. McGowran for his critical 
reading of the manuscript. Many of the con- 
cepts considered arose from discussions with 


ny colleague, Dr. M. R. Walter, now of Yale 
University. The figures were drafted by Mc. B. 
Frost, Department of Mines, and by my wife, 

The paper is published with the permission 
of the Director of Mines. 


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geal, Sac. §, Afr. 35, 29-36. 


Yours, R. B. (1933b) —Conditicns of deposition 
of the Dolomite Series. Trans. yweol. Sov, 5, 
Afr. 36, 191-135, 


Yours, R, B, (1935). A compurson of certain 
siramatolichic rocks in the Dolomite Series 
uf South Africa, with modern algal sediments 
in the Bahumus. Trans. peal. Soo, ¥. Afr. 37, 
153-167, 


Appendix 1.—Glossary 


Axis: The centre-line of a eoluoim, 


Bioheren) A circnmsctibed organo-sedimentary 

Structure whose minimum width is less than or 
¢qual to One hundred times ity maximum thick- 
ness, embedded in rocks of different lithology. 
Note: the definitions of the terms Aloherm und 
Biostrome pve based on those piven by Nefsan, 
Brown & Brineman (1962) but since, at Jeast 
in stromatolites, the two. integrade, an arbitury 
limit must be set. 
Tabular bieberms have parallel upper and lower 
surfaces, while deemed bioheems have gently 
convex upper surfaces. Subspherical hivherms 
had the highest growth relief relative to their 
width, 


Toasuing dbioherme are bioherms which had 
little or no growth relief, and therefore inter- 
tongic at their margins with the surrounding 
sediment. 


Blastveme: A stratiform organo-scdimentary arie- 
tire whose minimum width is more than one 
hundred times ity thickness, Note; jo practice if 
is rarely possible to see the Uiee dimensional 
shape of the strueture in outcrop. The distinction 
helween, bivherms and biostromites must there 
fore be based on the dimensions visible in out- 
crop. Ef the outcrop is inadequate, the informal 
tern) “hed” is used. 


Tabular biestremes have parallel lower and 
upper surfuces. Domed biestromes ether may 
cunsist of jietaposed domed bioherms or may 
be continuous with jirxtapased domes on their 
upper surfaces. 


Branching? The’ division of a column into new, 
diserete coltimmns. The columns became discrete 
when they are first separated by an interspace. 
In parallel branching. the aves of the new 
columns ure parallel (most commonly they are 
also parallel to the axis of the original column). 


a-puradlel branching is parallel branching in 
which the widih of the individual remains con- 
stant, Io a-paralle? branching the original column 
wklens geadually before branching, while in 
y-parallel branching, it widens abruptly before 
braaching, In sliphrly divergent branching. the 
nxes of the new columms diverge at less than 
45°. while in markedly divergent branchins 
they diverge al mure Man 45°. Dicharorioey 
branching (§ branching ips more than twer 
columns at approximately onc level, 


Bridge: A stronvatolitic lamina or set of laminne 
linking adjacent columns. 

Bump: A low, rounded pratrasion on the ade of a 
column. 


Coaleseing columas: Adjacent columns which joi 
und continue arowth as one column. 

Column; A discrete stromutolite structure, with the 
dimension in the direction of growth greater 
thum at least one of the transverse dimensions. 
Column shupe and arrangement offen vary ac- 
cording to the position in the bioterm, 

Columnur-layered strometelite: A stromatalite in 
which short columnar and laterally Unked 
fusually pseudocalumnar) portions ullernate. 

Cornice: Peripheral overhanging portion of a 
Jamina of set of laminae. elongated transversely 
to the column axis; 


Crest: The summit of an upward-convex Janina. 


Creéral line: The line jointing, the crests of succes- 
sive laminae 

Cresrat zene: The environs of the erestal line, In 
Conephyran, the crestal ome is specifically the 
yone of thickening and contortion of the 
laminae; the width of the crestal zone is the 
width of the thickened and/or contorted por- 
tions of Jaminac, Three types of crestal zones 
of Canophyion were distinguished by Komar 
et af. (1965), 


SOUTH AUSTRALIAN STROMATOLITES 93 


Cumiulate stromatelite: A rounded protruding nan- 
columnar stromatolile. 


Domed: With approximately constant radius of 
curvature, 


Flat-laminated strometolite: Non-cotumnar stro- 
mutolite with flnt continuovs laminae. Aitken 
(1987) has proposed the term cryplalgalaminate 
for stromatolites with planar lamination. 


Gently convex lamina, A, lamina whose ratio a 
hejght to diameter iy less than or equal to 0,3, 
Meusuremenis of this. ratio are best treated 


statistically by plotting on a histogram. 
Grarled columa: A column with large bomps 


Grumois: A mineral texture in which fine-grained 
Palehes ave surrounded by coarser grains, inter- 
preted jo have formed by partial reerystulliza- 
lion. 

Nypidiotopic: A mineral texture intermediate be- 
tween xenotopic and idiotopic. 

Miotapie: A texture in which the mineral grains 
ure hounded hy crystal faces. 


Individual: A single discrete stromatolite within 
which either the laminac are continuous or 
which comprises @ proup of columns arising 
from ia single basal column. 


Tiderspace: The space beuveen columns, usually 
filled with sediment. 


Lamina: The smallest unit of layering in a stroma- 
tolite. 


Lanceolare: An clangate Lransverse section of 4 
column. tapering at both ends, 


Laterally linked stromatalite: Stromatolite with 
wavy laminae which are continuous between 
crests. 4 


Mutralamina: A distinct set of laminae, 


Microstructure: Uhe fine-scale structure of the 
stromaltolite Jamination, in particular the dis- 
linciness, continuity, thickness and composition 
of the laminae. 


Banded microstrucnire is characterized by very 
continuens laminae with sharp, distinct, more or 
less parallel boundaries. In. streaky mticrosirue- 
ture less distinct and continuous laminae fre- 
guently grade into one another. The darker 
faminae are usually more distinct. 


Sttiated = micrystruciure consists of primary 
chains of Jonses, oriented parallel to the lamina. 
tion (this excludes cases where originally con- 
tinuous faminae are disrupted by recrystalliza- 
tion). 


Fermiform microstricture consists of narrow, 
Sinniows, pale calaured areas (uswally of sparry 


carbonate) surrounded by darker, usually finer 
grained areas. 

Micra-unconformity: Surface of lamination dis- 
cordance due to penecontemporancous erosion 
within a stromatolite. 

Niche; A deep indentation in the side of a colomn, 

Parabolic lamina; A lamina whose axial longitu- 
dinal section approximates a parabola. 

Peak: Overhanging portian of a lamina or set of 
laminae with a small dimension transverse to the 
column, 

Pellet; Ovoid to sub-ovoid micritic carbonate grain 
of silt or sand size, lacking internal structure 
Piginent: Organic or inorganic colouring matter. 
Plary celienn) A strongly transversely elongated 

eoluimn. 


Projection: A small columnar or conical out- 
erowth from the side of a column. 
Psenducelumnar: stramatolite: laterally linked 


stromatolite in which Successive crests are 
superimposed, forming calumn-like structures 
Cpsendocolumns), 

Rectangular lamina; Lamina which in a longitu- 
dinal section of a column js flal-topped with 
edges deflexed at about 90°. 

Rhombie lemina; Lamina which in a longitudinal 
section of a column is flat-topped but has sub- 
parallel edges not perpendicular to the top. 

Rib+ A low, rounded protrusion which ts elonguted 
transversely to the column on which il ozcurs, 

Sefvage: An unlaminated coating on colum mur- 
zing, Possible explanations for this include (a) 
micritization by algal boring: (b) inorganic pre- 
cipitation of lime; (c) a thin algal film an 
column margins during growth. In some forms 
4 selvageé-like structure is probably ihe result of 
differential recrystallization of a wall, 

Steeply Convex latina: A lamina whose ratio of 
height to diameter is greater than 0.5. 

Tuberous coliana, A coltimn with prominent ex- 
pansions and constrictions, 

Wil: Structure at the margin, of a column formed 
by one or more laminae from within the colunin 
bending down and coating the margin for at 
least a short distance 

Wavy lamina: A lamina with flesures of wave- 
length greater than 2 mm. 

Wrinkled lamina: A lamina with flexures of wave- 
length greater than 2 mm. 


Undulatory xtremurolite: Lateralty linked Stroma- 


tolite in which successive crests are not super- 
imposed, 


Xenolapic texture: A texture in which the mineral 


grains are anhedral or irregularly shaped, ie. 
net bounded by crystal faces, 


4 


Fig. 10, 


Fig. 11, 


Figs 12 


Fig, 13. 


Pig. 14. 


Fig. 15, 


W. ¥. PREISS 


Acaivila angepena, Irom Lower Cambrian limestones, Flinders Ranges: sections perpendicu- 
liar to bedding. showing mode of occurrence and microstructures; (a) —Marginal section of 
a bioherm, Note thut the laminae are completely recurved wnver the biohetm evige. The speci- 
men is in sifu. Ube ball-point pen is 16 cm Tong, Angepena; (b)—Etched section of Sab, the 
recurved margin of the bioherm in (a) cur at right angles (o bedding, Nate that here growth 
partly proceeded downwards, Specimen is 15 cm wide; it was collected from the surtcrop 
shown in (ay; (co) —Laleral termination of a bioherm, which partly sank into the soft substrate 
during growth, The white areas are dolomitized. Width of specimen (S460) is 20 cm. Ange- 
pent; (d)—Pseudocolumns with rare interspuces, Note the domed Liminae grown upon partly 
Kiril intvaclasts, and the extremely continuaus lamination, Uhin section. Angepena (8462); 
(ej—Evenly laminated ferruginous structure, probably the stromatolite A. aneepena affected 
by secondary ferruginization, Thin section, from near Old Wirrealpa, The durk laminae are 
outlined by finely disseminated hematile. (S364, collected by Mr. P. G. Haslett), (f)—Dyenly 
lanimated discrete columnar form from halowie Gorge (Sir Douglas Mawson's specimen), 


(a j—Arociely dagepenc; irregular coluoins freny ihe marginal portion of @ small bioherm 
15458 Anwepena), Note the vermiform microstructure within parts of columns, here inter- 
reted us duc to algal boring, disrupting the normally very even, continuous lamination; (b) 
» de)—aAeaciella f, indet, Both specimens are erraticy from the Storiian glacials north of the 
Enorama Dispir. Thin seaions. Note the very numerous concordant stylolites in (bh). (S539 
and $509 respectively; S$39 was collected by Dr. B. Daily); (dl. Ce) & CE) —Aceetella 
wngustu, Brighton Limestone equivalent, Depot Creek, Vertical sections showing mode of 
Ocuurrence aml microstructure; (dJ—-Details of transition from broad, frequently bridged 
basal colunmms to upper nurrow, discrete columns. Broad coltimns in lower right-hand corner 
five inclined margins and subhoriznntal laminae: tel--Lenticular open spaces between 
uminue, possibly representing origina) gay vesicles (S163). (f)—Portion of a bioherm show- 
ing the inlercalytion of columnar snd laterally linked siromatolites. 


Avueiella wigusta, Brighton Limestone: equivalent, Depat Creek, showing mode of wocurretice 
and ticrosiructures. (9)—Margin of a bioherm (pale colonsed at right of photograph) inter. 
tonguing laterally with massive ooypuciie (at left); (hb) & (c)—S4U4 & S401 respectively. The 
gross shape and branching of columns, The interspuces ure filled wit inteclayered micrite and 
intramicrite, In U5 (o L.Weem bands. te) Is natural size. In (b), laminae become daubly 
Crestcd before branching, but in the centre of the photograph (c) is an example of a short 
interspace beiween crests bridged by the overlying laminae: the voltimn then resumes ils former 
growth pattern; (d)—Recryslallized specimen from Mundallio Creek (3538). illustrating ra- 
isting recrystallized acicvlor textiires im the Lower part of the pholagraph. 


Ralrwia Narra, Skillogalee Dolomite; Sections perpendicular to bedding. showing the mode of 
focurrence and microstructure: {a)—Small lenticular biohermes interbedded in thinly bedded 
dolomites, Yalina; (b)—Portion of a biostrome interbedded in massive, fine eiained dolo- 
mites, Dutton’s Trough H.S. Longitudinal section of partially: silicified columns. The section 
is parallel to the tectonic cleavage, in the plane of Mattening of the columns; (c)—trregular 
eolumnas with numerous mickruncontormitics and highly variable lamina shape, West Mount 
Hur td)—Moderarely divergent branching columns, with some pelletsl laminde, Thin séc- 
tion, Yutina ($222, holotype); (c)—Stightly divergent branching in regilar, sub-cylindrieal 
collimns. Thin section, $533, Duiton's Trough HS, the specimen is taken from {he biostrome 
shown in Fig. 13 {h)} 


Reivwlia barra, Skillogalee Dolomite: (a)—Tuberouvs and inclined columns with evenly 
handed microstructare and high-angle micro-unconformities, Thin section, S487, Myrtle 
Springs; (b) BR. barra with minor pelleial laminae. Thin section, S150, Worumba, (c)—Sub- 
evlindrics) columns with steeply domed, evenly banded laminae, Thin section, $302, West 
Mount Hut: (d3—Branching of narrow columns from the sides of a main wide column, Cut 
slab, 5534, Dutton’s Trough H.S. The specimen is taken from the blastrame shown in Fig. 
13 (6b): fev}—#, burre with predominantly pellets! luminue. Thin section, $496, Copley: (EJ 
—Conplex branching of columns [ram Arkaroola, Thin section, $457. 


(aj—Baicalio Suera with finely silicified laminue. Thin section, natural size, S151, Warumba. 
Note the vertical tectonic dolomite veins; (b)—#. burra. Cut slob tliustrating sub-parallel 
bronehing columns with high-angle micro-unconformilies and bunded tamination. 396, near 
Chintapanna Well. specimen collected by Mr. C. R. Dalgarno, Note the overgrown siroma- 
tolite fragment in the lower left quadrant, and he branch arising from an_eroded column ih 
the Upper right; fe}—Indcterminate siromatolite, possibly Baroulia barra, Thin section, $322, 
near Cartieion: (4), fe) & (£)—Boxonia melrose, Brighton Limestone equivalent, Melrose: 
fd)—Hand specimen iflustrating longitudinal sections of columns: (¢)—Thin section of halo- 
type. 5503, Che lamination is indistinctly banded, and becomes diffuse in the wall zone, (f) 
—Thin section illustrating Jamination and wall structure, S177, natural size, Note that the 
upper left and lower left corners of the thin section ure composed ot highly weathered rock, 


YS 


SOUTH AUSTRALIAN STROMATOLITES 


10 


“1G, 


"6 W. V. PREISS 


| 7 PON 


™. 
ae 


a She SAAS 
a ; 


FIG. 11 


97 


SOUTH AUSTRALIAN STROMATOLITES 


12 


FIG. 


98 


W. V. PREISS 


SOUTH AUSTRALIAN STROMATOLITES 94 


FIG. 14 


PRETSS 


wv 


100) 


abe te 


x 
Lj 


“a 


THE NEMATODE GENUS MAXVACHONZA (OXYURATA: 
COSMOCERCIDAE) IN AUSTRALIAN REPTILES AND FROGS 


BY PATRICIA M. MAWSON 


Summary 


The genus Maxvachonia Chabaud & Brygoo, 1960, previously known only from reptiles in 
Madagascar, is now recorded in Australia and New Guinea. New species described are M. chabaudi 
from 7 species of skinks, 1 species of gecko, and 1 species of snake (? from food); M. brygooi from 
5 species of agamid lizards; and M. ewersi from a frog, Litoria nasuta, from New Guinea. M. 
ftindersi (Johnston & Mawson) [syn. Aplectana flindersi J. & M.], 1s recorded from 5 species of 
Australian frogs and one introduced species, Bufo marinus. The genus Austrocerca Inglis, 1968, is 
regarded as a synonym of Maxvachonia. 


THE NEMATODE GENUS MAXVACHONIA (OXYURATA: COSMOCERCIDAE) 
IN AUSTRALIAN REPTILES AND FROGS 


by Patricia M, Mawson* 


Summary 


The gétnus Maxvachonia Chabaud & Brygoo, 1960, previously kaown only from reptiles in Mada- 
Rascar, 8 now recorded in Australia and New Guinea. New species described are M. chabauedj Irom 
7 species of skinks, | species of gecko, and 1 specics of snake (7 from food); M, brygeoi from 5 
species of agamid lizards; and M. ewersi from a Frog. Litoria nasuta, from New Guinea. M_ flincersi 
(Johnston & Mawsun) [syn. Apleetana flinderst J. & M,), is recorded from 5 species of Australian frogs 
and -one introduced species, Bufo marines. Che genus Anstrocerca Inglis, (968, is regarded as a syno- 


nyin of Maxvachania. 


Introduction 

Maxvuchonia dimorphe Chabaud & Brygov 
(1960, p. 129) was first described from 
Chamaeleon pardalis, and tater also from C. 
australe (Chabaud, G. R. Caballero, & Brygao 
1964, p. 846}, in both cases from iw small 
island. Nossi-Bé, about 20 km from the main- 
land of Madagascar. It has since been recorded 
from one chameleon and two otler specics of 
hzards (Zonesauras maxinus and Mebuie 
gravenhorviii) on Madagascar itself. (G. 
Cuballero 1963, p. 192.) 


Although the genus was net recognised until 
recently, and docs not appear to be common 
in any host species, it is surprisingly wide- 
spread. The seventeen species of Australian 
lizards from which Maxvachonia spp. are 
recorded in this paper belong to the families 
Scinctdae, Agamidae, and Gekkonidae, and 
they come from a wide geographical range. One 
collection was. made from the stomach of 4 
snake. but as this also contained some semi- 
digested skinks, the snuke may not be a true 
host record. 


The gents is not confined to reptiles, A ustra- 
cercu Inghs (196%, p. 164) appears to be a 
synonym of Muxvachonia. Inglis recorded 4. 
finders? (lotision & Mawson) (syn. A plectana 
finders!) from three frog species in Western 
Australia, Tt has now been recognised from five 
mute frog species from various parts of Aus- 
tralia, and from a twad, Bufo marirtus, intro- 
duce? inte Queensland sugar cane fields in 
1934, Another species is recorded from a frog 
from New Guinea, 


The males and females of Marvachenia spp. 
we very different in size, but the morphology 


of the anterior end is similar in the two sexes. 
Both males and females are easily distinguished 
from other cosmocercoid genera, the Female by 
the great distance of the anus from the pos- 
terior end of the body, znd by the shape of the 
eggs, und the tale by the shape of the guber- 
nuculum, which is very large and bears two 
prominent projections near its proximal end. 

The differentiation of species within the 
genus is rather more difficult. The presence or 
absence of lateta] alae on the anterior part of 
the body in the female appears to be a specific 
character. There is a wide varislion in the body 
length of the female within a species, although 
fully adult specimens from the same host ant- 
mal are usually about the sanic size, The ratio 
of the body length to that of the oesophagus 
varies considerably, possibly due at Jeast in part 
to the degree of contraction of the body in 
different collections. The ratio of body to tail 
length in the adult is more constant, and may 
be of specific significance. The em size is simi- 
lar in all specimens available, but there is some 
vanation in the shape of the projection on the 
egg shell and of the envelope which surrounds 
the egg in the vagina, and these appear to have 
specific. value. 


The male worms are rare compared with the 
female. so thal it is even harder to assess the 
specific value of any character. The body 
length and that of the ocsophagus are very 
similar among all the specimens examined. 
‘There is some vuarigtion in the iengths of spi- 
cules and gubernaculum but even these vary 
almost as much between two specimens from 
the same host animal (in the only case where 
two males were found in one host) as among 
all the males collected in Australia. 


* Zoology Department. University of Adélaide. Adelaide, S$. Aust. 5000. 


Truns, R. Soc, S. Aust. Vol, 96, Part 2, 3! May 1972, 


1bz PATRICIA. M. MAWSON 


On theve slender ctiterin, three species have 
heen distinguished from Australian reptiles, one 
(M. flinderné) fron Australian frogs, and onc 
from a New Guinea frog. The general body 
form is similar in all species, and agrees gene- 
rally with the descriptions of Chabaud & 
Brygoo (1960) and Chabaud et al (1964) and 
o£ Inglis (1968), Some additionul observations 
and westinguishing characters are noted under 
the species. Measurements are given in Tables 
lanst Hl. Pype specimens will be deposited in 
(he South Austtalian Museum, 


Maxvachunia chabaudi nap. 
FIGS. |-6 


Hosts and lucalities: Murerhia 
{(Dumerit & Bibron}, iwpe host, Leriva 
hoacaimillii (Gray), Crenoras — leae 
(Boulenger), Psendonaja ? affints Ciiinther, 
all from Eyre Peninsula, S. Aust.; Crenofus 
labillurdlevi (Gray) from Pemberton, W. 
Aust; Afeviergis peronil (Fitzinger) from 
Pemberton and Esperance, W. Aust.; Spivrte- 
niorplius chxtralis (Gray) from Wilgarup, W. 
Ausut S. koscinsko’ (Kinghorn) from the 
New England district, MUS.W.; Egernia 
whirel (Lacépie) from Penola, S. Austy 
Phyllurus silit (Bory) from Kangaroo 1), 3. 
Aust, 


Must of these collections consist of female 
worms. adult and/or juvenile (i.e. with or 
Without embryonated eges). There arc five 
males, two from Crenotux leae, one from 
Leriste lineoecellatuy and one in cach of two 
H. pevonii, Tn these last two there were no 
Temules, and as this species of Maxvuchonic is 
separated from others by chiuructers of the 
female. the inclusion of the males js arbitrary. 


Alac ure present in both sexes. There are 
three lips, the inner border of each projecting 
as a cuticular lamella. The mouth is triangular, 
or triradiate, Euch Jip is strengthened by it 
chitinous bar, the three bars mecting to form 
a triangle around the anterior end of the buccal 
cavity, The short triungular buecal capsule 
rests against the anterior end of the eesophagus 
Three well-defined teeth project fram the oeso- 
phageul lining into a depression in the anterior 
end of the oesophagus. 

Female: Lateran] alge extend from the level of 
the nerve ring to about the mid body. The pos- 
terior end of the hody ends in a more or less 
distinct mucro, which is rugose, The vulva, a 
transverse slit, lies at about the level of the 
isthmus of the oesophugos. The two ovartes 
commence shortly in front of the anus, pass 


lineoocellara 


backwards nearly to the posterior end of the 
body where each enters a short oviduct, lead- 
ing to a slightly wider, sometimes almosr 
spherical, thicker-walled sectiun {? seminal 
receptacle) from which the uterus Ieuds for- 
ward. The two uteri pass forward sido by side. 
uniting to form the vagina at about » quarter 
of the body Jenpih Lrom the anterior end, or a 
little in front of this. 

Eggs in the anterior parts of the uteri cach 
contain a coiled Jarva The eyes are rouchly 
sphericul, slightly longer in the axis through the 
knob on the shell, In the vagina, where they are 
less crowded, they ire seen to be surrounded 
by 4 spongy or reticulate material which Lorms 
a loose envelope attached to the shell by, or iat, 
the apica) knob, mare or less open at the oppu- 
site pole (Fig, 5) «ind often trailing two ribbon- 
like pieces from the open end. This envelope 
was noted in the ariginal description of Mutva- 
chania dimorpha, 


Male: The lateral alae extend for most of the 
body length, from the Jevel of mid-oesophagus 
to shurily in front of the anus, The posterior 
end of the body is strongly curved ventrally. 
The gubernaculum is Jarge and heavily built, 
With q pair of lateral processes neat the proxi- 
mal end. The spicules are slender. well chiti- 
nised, sind blunt-tipped. The cloacal opening is 
on un clevation of the body wall. ‘The thirteen 
pairs of caudal papillac are arranged as shown 
in Pig. 6. 

The species is distinyuished irom M. di- 
morphea chiefly because of the presence of 
lateral alae in the female. The females ave all 
shorter, and the males about the same size, as 
those of ML dimaerpha bot the spicules and 
gttbernacuilum are larger, 


Muavachonia brygoot nap, 
PIGS, 7-10 
Hosts ind localities: Amphibolurus. decresti 
(Dumeénil & Bibron), type bust; A. nracrelarns 
(Gray), both from Eyre Peninsula, S. Aust.; 
A. inermis (De Vis) izam Yuendunu, North- 
erm Territory; 4, muricatus (Shaw) and A, 
herbotay (Cuvier) from N-S.W~ 
Only femples have heen taken from these 
agamid lizards. All of them, however, differ 
from those from skinks in the absence of lateral 
alae. In other respects they are very similar. 
Althotigh this distinction is slight, it is con- 
stant Notwithstanding the fact that agamids 
and skinks aceurret in the same tocality in 
Hincks National Park an Eyre Peninsula, Afiex- 
vachowa spp. from the agamids were always 


Figs. 1-6. 


MAXVACHONIA IN AUSTRALIAN REPTILES AND FROGS 103 


Maxvachonia chabaudi. Fig. 1.—Oesophageal region. Figs. 2, 3.—Lateral and en face 
views of head. to same scale. Fig, 4.—Posterior end of femalc. Fig. 5.—Egg. Fig. 6— 
Posterior end of male. 


104 PATRICIA M. MAWSON 


without alae, while those from the skinks had 
alac. In view of this it is thought safer to regard 
the two groups as separate spccics, at least until 
more specimens, especially males, are found. 
Maxyachonia sp, 

Host and locality: Morethia taeniopleura, 

Mornington I, Gulf of Carpentaria. 

Only one female was collected from this 
host; it is very similar to females of M. 
chabaudi but the ratios of oesophagus and tail 
{o the body length differ markedly (Table 1). 


Maxvachonia filndersi (Johnson & Mawson) 
FIGS. 11-13 
Aplectana flindersi Johnston & Mawson, 
1941: 148, from Liroria ewingi (syn. Ayla 
jervisensix) from Kangaroo L, 8. Aust. 


Figs. 7-10. 
end of female. Fig. 10.—Egg. 


200. um 


Ausirocerca flindersi (Jobnston & Mawson) 
Inglis, 1968: 165, from Literia coelo- 
rhyncha, Heleioporus barycrugus and A. 
psrammophilus, from W. Aust. 

Host and localities: Bujo marinus Linn. from 
Queensland: [.imnedynastes dorsalis (Gray) 
from Adelaide, $. Aust.; Heleioporus ittor- 
naatus Lee & Main, Litoria moorei Copland, 
L. adelaidensis. (Gtay) from near Perth, W. 
Aust.; L. caerulea (White) from Alice 
Springs, N.T. 

All the hosts listed above are new records 
for M. flindersi. The new male specimens 
agree closely with the earlier descriptions, both 
in size and appearance, but the females are dis- 
tinctly larger; even those from related hosts in 
Western Australia. Through the courtesy of 


Maxvachonia brygooi. Figs. 7, 8.—Lateral and en face views of head. Fig. 9—Posterior 


Figs. 11-13. M. flindersi. Fig. 11—Anterior end of female, Fig, 12.—Posterior end of female. Fig, 


13.—Egg. 


MAXVACHIONIA IN AUSTRALIAN REPTILES AND FROGS LOS 


Dr, W. G. Inglis and of the Western Australian 
Museum it has been possible to compare all 
the known specimens, and no significant differ- 
ence other than size was observed. The details 
of the female reproductive system have now 
been studied, and these agree generally with the 
form in other species of the genus, The ovaries 
begin shortly in front of the anus, The eggs in 
the uteri are enclosed in the characteristic outer 
envelope, which in some specimens is very 
dark. The envelope is in the form of a bell 
atlached to the knob of the shell at its apex 
and open at the other end; from the open end 
come two long ribbons of material similar to 
that of the envelope. In one case an egg lying 
fast outside the body of the female was still 
attached by one of these ribbons, which passed 
into the vulva. 

Maxvachonia flindersi differs from MM. di- 
morpha in the presence of well developed 
lateral alae in the female, and from both M. 
dimorpha and M, ewingi (see below) in the 
shape of the seminal receptacle. The size of the 
spicules and gubernaculum vary greatly in the 
few male specimens known, but the gubernacu- 
Jum is always distinctly longer than the spicules, 


14 


BOG LTT 


E 
a 
I9s 


60 Lin 


Maxyachonia ewersi 1.sp. 
FIGS, 14-20 
Host and locality: Lireria nasuta (Gray) from 

Brown River, New Guinea. 

The material consists of three female and 
one male worms. The genera! body form is very 
similar to that of M. flindersi and other species 
of the genus; measurements are given in Table 
Hs 


The churacters distinguishing this species 

from M. flindersi are the following: 

1. The oesophageal teeth are much smaller 
(Fig. 15). 

2. The spicules are distinctly longer than the 
gubernaculum. 

3. There are only two pairs of preanal papil- 
lae in the male. The other caudal papillae 
are arranged as in M. flindersi, 


4. In the female the posterior end of the hody 
uppears rounded, becuuse the extreme tip 
is slightly withdrawn forming a dimple. 

5, ‘The artaiigement of the reproductive organs 
in the female is slightly different. The 
ovaries start much further forward at about 


Pigs. 14.20, Maxvachonia ewersi. Fig. 14.—Oesophageal region, female. Fig. 15.—Lateral view of 


head, Fig. 16—Posterior end of female, Fig, 


{7.—Fge. Figs. 18, 19—Two views of 


apical extension of egy shell, to same scale, Fig. 20.—Posterior end of male. o. ovary; 


Tr, seminal receptacle; u, uterus, 


106 


two-thirds the body length from the head, 
and the seminal receptacle is not so much 
wider than the uterus and ovejector, 


6. The shape and size of the eggs are different. 
The knob on the shell is shorter, and 
appears conical on one axis, but hroad and 
grooved on un axis at right angles to this; 
the egg itself is slightly flattened in this 
latter view. The coils of the larva lie in the 


PATRICIA M. MAWSON 


plane of the wider diameter. The envelope 
surrounding the egg is thinner than in other 
species, although it is dark in colour, and 
forms a bell, attached at knob end of the 
egg, similar to those of M. flindersi, but 
more definite in shape (in these specimens 
at Icast). In eggs furthest from the vulva 
(bul in the vagina) the mouth of the bell is 
open, but in those nearest to the vulva it is 
closed. 


Acknowledgements 


Several of the collections examined were 
made by Dr. John Hickman of the Zoology 
Department, University of Tasmania; speci- 
mens from Literia nasuta were sent by Dr, W. 
Ewers of the University of Papua and New 
Guinea. To both these helpers I am most grate- 
ful. 1 also wish to thank Dr, W, G. Inglis, then 
Direcior of the South Australian Museum, 
because I would not have examined. any frog 


material for Maxvachonia sp. had he not 
pointed out that his genus Austrucerca is 2 
synonym of Maxvachonia. 


I am also indebted to officers of the South 
Australian Museum, Mr. M. Tyler, Hon. Asso- 
ciate in Herpetology, and Dr, T. Houston, 
Curator of Amphibia and Reptiles, for infor- 
mation on the nomenclature of the hosts. 


References 


CABALLERO, G. R. (1968)—Contributions @ la 
connaissance des nématodes da sauriens Mal- 
gaches. Annis. Parasit hum. comp. 43, 149- 
200. 

CHazaup, A. E. & Bryaoo, EB. R. (1960). Neéma- 
todes parasites de caméléons malgaches. IL 
Mem, Inst. Scient. Madagascar A. 14, 125- 
159, 

Cuasaup, A, E., & BryGoo, E. R. (1962)—Néma- 
todes patasites de caméléons malgaches. 
Deuxiéme Note. Annis. Parasit. hum. comp. 
37, 569-602. 


Cuapaup, A. C., CABALLERO, G, R., & Brycoo, 
E. R. (1964).—Affinilés entre les genres 
Skrjabinelazia (Ascaridida Seuratoidea) et 
Maxvachonia (Ascaridida Cosmocercoidea). 
Bull, Mas, Nat. Hist. Nar., Patis 36, 844-848. 


Incuis, W. G. (1968)—Nematodes parasitic in 
Wester Australian frogs. Bull. Brit. Mus. nat. 
Hist. Zoology. 16, 163-183. 


Jounston, T. H., & Mawson, P. M. (1941).— 
Some nematodes from Kangaroo Island, S.A. 
Ree. S. Aust. Mus. 7, 146-148. 


107 


MAXVACHONIA IN AUSTRALIAN REPTILES AND FROGS 


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THREE NEW SPECIES OF THE GENUS CLOACINA LINSTOW 
(NEMATODA: STRONGYLATA) FROM MACROPOD MARSUPIALS 


BY PATRICIA M. MAWSON 


Summary 


Three new species of Cloacina are described: C. mundayi from Macropus rufogriseus, from 
Tarraleah, Tasmania, characterised by the presence of a dorsal buccal tooth associated with the duct 
of the dorsal oesophageal gland; C. clarkae from M. eugenii, from Kangaroo Island, South 
Australia, characterised by the shape of the cephalic papillae and the structure of the oesophagus; C. 
edward,vi from M. bicolor, from Sunday 1, Victoria, characterised by the presence of oesophageal 
‘plumes’, very short spicules and very short vagina. 


THREE NEW SPECIES OF THE GENUS CLOACINA LINSTUW 
(NEMATODA: STRONGYLATA) FROM MACROPOD MARSUPIALS 


by Patricla M, Mawson” 


Summary 


Three new species of Clog¢ina are described: ©. 


miadayt front Macrepus rifegrivens, Trom 


Vactaleah, Tasmania, characterised by the presence of a dorsal buccal tooth associated with the duct 
of the dorsul oesophageal gland: tC. clarkae from M. eugenii, from Kangaroo Island, South Australia, 
characterised by the shape of the cephalic papillae and the structure of the oesophagus; C. edwardsi 
from Af, hicéfeor, from Sunday U., Victoria, characterised by lhe presence of oesophageal ‘plumes’, very 


short spicules and very short vagina. 


Cloacina mundayi n.sp. 
FIGS. 1-7 


Host and locality: Macropus rufogrised, from 

‘Varcaleah, Tus. 

This species is u relutively short stout nema- 
lode. The material consists of four males and 
four females. The submedian papillae are small, 
with the distal segment much smaller than the 
proximal. The long threadlike cercival papillae 
are close to the anterior end. 

The shallow buccal ring is somewhat hexa- 
gonal in shape, and is uneven in depth, A snl! 
dorsal ocsophazeal tooth projects into the 
buccal cavity, and is traversed by a duct from 
the dorsal oosophageal gland, The oesophagus 
is cylindrical, widening only slightly at the pos- 
Tertor und. No teeth ure present in the lumen. 
The nerve ring is at about the mid-length of 
the oesophagus, and the excretory pore shortly 
behing it. 

The posterior end of the female tapers from 
in front of the vulva to the tip of the tail; the 
distance from the anus to the vulya is ubout 
equil to the tail length, The yiagina is slightly 
longer than the tail, The eggs are 110 by 65 
phy 

The spicules aré ahout 1/2.5—-3,0 of the bady 
lcngth.. The dorsal lobe of the bursa is unusu- 
ally Jong for the genus. and the ventral lobes 
are united, The bursal rays are as shown in 
Figs. 6 and 7, The genital cone is short and 
conical, and on either side of it there is a 
cuticuJar mflation. No accessory cone can be 
seen, Measurements are given in Table 1, 

This species is distinguished from all others 
so far described in the shape of the bursa and 
in the presence of a dorsal tooth in the buccal 


capsule. In describing C. dahl’, Linstow (1897, 
p. 287) mentions the presence of a gland (seen 
in T\S,) in the dorsal wall of the oesophagus, 
with «a duct opening dorsally (presumably into 
the lumen of the oesophagus or into the 
mouth). Such a distinct gland has not been 
noted in descriptions of other species of the 
genus, nor in re-examination of fresh material 
of various species, Tl is not present in €, 
clarkue ov CG. edwerdsi. In the deseription of 
©. daidi there ix no indication of a dorsal touth 
associated with the gland. 


Cloacina clarkae nsp- 
FIGS, 8-13 


Host and locality; Muacropuy eudgenii, from 
Kangarou T 


This is a large worm from the stomach of 
the host. The submedian papillae are Jong and 
slender, and the distal segment of euch is dis- 
tinctly longer, but not wider, than the proximal 
one. The cuticle is thickened just behind the 
cervical grouve, The threudlike cervical papillae 
are Telatively close to the anterior end, The 
buecul ring is deep, its walls relatively. thin, and 
sloping outwards towards the anterior end; the 
unteriar muryin is lobed. 


The oesophagus is Jong and slender, except 
for a distinct terminal bulb, It is clearly divided 
mio four remions—(1) in the anterior half 
there are 9-11 distinctive equidistant places 
where the lining appears to he creased; (2) in 
the rest of the cylindrical part of the oeso- 
phagus the cuticle is more or less featureless; 
(3) just below the terminal bulb, the oeso- 
phagus and its lumen are slightly wider, and in 
this region about 8 well deyeloped teeth project 


* Deparment of Zoology, University of Adelaide. Adelaide. S. Aust. 5000. 


‘Trans. R. Soe. 8. Aust. Vol. 96, Part 2, 31 May 1972. 


110 PATRICIA M, MAWSON 


50 


j 


Figs, 1-7, Cloacina mundayi. Fig. 1.—Oesophageal region. Fig. 2.—Head, lateral view. Fig. 3— 
Head, en face, Fig. 4—T.S. anterior end shortly behind buccal ring. Fig, 5.—Posteriar 
end of female. Figs. 6. 7—lateral and ventral views. of bursa. 


Figs. 8-13. Cloacina clarkae, Fig. 8—Lateral yiew of oesophageal region. Fig. 9- Head, ventral 
views. Fig. 10.—Part of oesophagus showing teeth in tumen. Fig. 1].—1],ateral view of 
bursa. Fig. 12.—Dorsal ray and genital cone. Fig. 13.—Posterior end of female. 

Figs, 14-22. Cleacina edwardsi. Figs. 14, 15—Lateral and en face views of head. Figs. 16, 17. 
Oesophageal regions of male and female respectively. Fig. 18.—Part of oesophagus shaw- 
ing plumose structures, Fig, 19,—T,.S. oesophagus in region of plumose structures. Fig. 
20,—Posterior end of female. Figs. 21, 22—Dorsal and laieral views of bursa. 


CLOACINA FROM MACROPOD MARSUPIALS iT] 


Figs. 8. 13, 16, 17 and 20 to scale beside Fig. 13. Figs. 14, [5, 18 and 19 to scale beside Fig. 19. 
Figs. 9 and 1() to scale beside Fig. 10. Figs. 21 and 22 to scule beside Fig. 22. 


We PATRICLA M. MAWSON 


into the fhumen (Figs. 8, 10); (4) the terminal 
both. Vhe nerve ring lies at about a third the 
length of the oesophagus from the head, and 
the excretory pore wt the level of the anterior 
end of the oesophageal bulb. 


In the feniale the tail is conical and pointed, 
and the yulva is about half the tail Jongth in 
fron! of the ants. The yagina is rather longer 
than the distance from the vulva to the tip of 
the tail, and is somewhat convoluted, The eggs 
are about 173 x 30 am, and contain a “tad- 
pole’ stage larva, 


The bursa ts much shorter ventrally than 
dorsally. The arrangement of the rays is shown 
in Figs, |) amd 12. The genital cone is well 
deycloped and bears dorsally a short pair of 
appendanges, The spicules are a little more than 
a quarter of the body length: a gubernaculum 
is present. 

The species is among the medium-large sized 
Cloacina spp.. and can be distinguished by the 
characters of the head and ocsophagus. It is 
perhaps closest to C. communiy Johnston & 
Mawson (1939), which however ts larger, and 
in which the oesophageal teeth are arranged 
ditterently (Mawson 1961, p, 196). 

The specific mame is given in recognition of 
the work of Miss Helen Clark who tsolated the 
worm, and who included a study of the eurly 
stages af the life history In work for an 
Honours Degree in this Depnriment. 


Cloacina edwardsi asp, 
FIGS, 14-22 
Hast and locality: Wallibia bicolor, trom Sun- 
day '. Vie 
This apparently new species of the genus 
Cloncina belongs to the group in which the dis- 
tal segments of the submedian cephalic paptilae 


are much larger than the proximal segments. 
The cuticle behind the head is thick, becoming 
less su towards the base of the gesophagus, The 
long threadlike cervical papillae lie shortly in 
front of the level of the nerve ring. The buceal 
ting is short, wide and stoutly built, the walls 
gyal to triangular in section, 

The oesophagus is short, more or less cylin- 
drical, with a small terminal swelling in the 
female, but not in the male. ‘There are no teeth 
in the lumen, but there are three very distinct 
“nlumose"” areus, one on each of the three walls 
of the lumen (Figs. 18. 19). in the region just 
unterior to the nerve ting. These areus arc 
formed by confluent ridges on the cuticle lining 
the himen. ‘They appear to be similar in form 
to such structures figured and described for 
some Marshidia spp. from elephants and rhina- 
ceros, but have not previously been described 
from Australian trichonematines, 


The nerve ring surrounds the o¢sophagus al 
or just behind its midlength. and the excretory 
pore is jist post-oesophageal. 

The posterior end of the femule tapers gra- 
dually from about the vulva, enditg in i slen- 
der pointed tail; the vulva is tather more than 
a tail length in front of the anus. The vagina 
is very short, The eggs are 83 4 50 aim. 

The spicules are Unusually short for Cloacini 
spp., about 1/14 of the body length; a guber- 
miculum is. present. The genital cone is well 
developed, conical, with two small projections 
forming the accessory cone. The form and 
arrangement of the bursal rays are shown in 
Figs, 21 and 22. 

This species is. distinguished from any pre- 
viously described by the presence of the plu- 
mose structures in the Gesophagus, as well as by 
the unusually short spicules and the very shorl 
vagina. 


Acknowledgements 


Lam very much indebted to the people after 
whom the species are named, who. have col- 
lected These and other nematodes and sent them 
to me—Mre. Barry Munday of the Mt. Pleasant 
Luboratories of the Tasmanian Department of 


Agriculture, Launceston, Tasmania, Miss Helén 
Clark of Adelaide, and Mr. Geoff Edwards. a 
post araduate student (1969) in the Depart- 
ment of Zoology, Monash University, Vic- 
torin. 


References 


Jounsvon, T. HL. & Mawson, P. M_ (1939) — 
Strongyle nematodes from Central Australian 
kangaroos and wallabies, Trens, R- Sor, S- 
Anat, G2, 263-28, 

Linstow, QO. vow (1897),—Nemuathelminther 
gesummele von Herm Praf. Dr. F, Dahl im 


Bismarck-Archipel. archiy, & Marturges. 63, 
281-291, 

Mawson, P. M. (1961).—A note on the occur- 
rence of oesaphogeal iceth in species of the 
seniis Cfloatina (Nematoda: Strongyloides), 
Trans, R, Soc, S$, Aust, 85, 85-89. 


CLOACINA FROM MACROPOD MARSUPIALS 113 


TABLE 1 


Measurements of Cloacina edwardsi, C. mundayi and C. clarkae; unless otherwise indicated, measure- 
ments are in um 


C. edwardsi C. mundayi C. clarkae 
male female male female male female 

Length (mm) 6.3 6.5—7.7 3.1-3.4 4.45.0 8.8-9.2 12.3-13.6 
Oesoph. length 480 470-520 470-490 470-570 1100-1220 1250-1350 
Antr. end—nerve ring 260 250-290 250-260 250-320 300-350 300-350 

—cervical pap. 220 170-200 100 80-90 115-160 110 (3x) 

—excret. pore 550 480-610 250-340 330-350 890-900 930-1020 
Spicule length 430 —_ 1100-1150 — 2400-2600 _ 
Tail length — 290-300 —_— 210-250 — 260-350 
Vulva postr. end — 680-730 —_— 410-500 — 430-500 
Length/oesoph. L 13 13-15 6.5-7.1 8.6-9.3 7 2-8.5 9,5—-10.7 


Length/spic. L 14.6 — 2.83.0 = 3.43.7 — 


FURTHER RECORDS OF THE PITTED-SHELLED TURTLE 
(CARETTOCHELYS INSCULPTA) FROM AUSTRALIA 


BY R. SCHODDE, I. MASON AND T. O. WOLFE 


Summary 


Further records of the Pitted-shelled Turtle, Carettochelys insculpta, including the first breeding 
record, are reported from northern Australia. It is concluded that the species occurs in river systems 
right around the landward margins of the Sahul Shelf. The stone fruits of Pandanus, whenever they 
are available, appear to comprise an important item of the turtle's diet. 


FURTHER RECORDS OF THE PITTED-SHELLED TURTLE 
(CARETTOCHELYS INSCULPTA) FROM AUSTRALIA 


by R. Scuoppr,* L Mason,* and 'T. O. WoLre* 


Summary 


firther records of the Pitled-shelled Turtle, Caretlechelys inseulpia. including the first breeding 
record, ace reported from northern Australia, 1 is concluded that the Species occurs im river systems 
vight around the landward margins of the Sahul Shelf. The stone fruits of Pandanus, whenever They 
are available. appear ja comprise an important ilem of the lurtle’s diet. 


The Pitted-shelled Turtle! Caresoehelys 
insclilpre Ramsay, sole living species oF a 
family, Curettochelyidae, that apparently 
vecurred widely in palaearctic and nearctic 
revions up until early Tertiary times, was 
known only from the river systems of southern 
New Guinea until 1969 (de Reoij 1915, 1922: 
Wermuth & Mertens [961). In that year, the 
first specimens were recorded for Australia 
(Cozver 1970). Ten specimens were captured. 
all in the Daly River, Northern Territory, in 
freshwater teaches about 13 km above tidal 
influence. Because the single specimen exa- 
mined did not appear to differ significantly 
from New Guinean specimens, and na evi- 
dence of breeding was found, Cogger (\.c.) 
speculated on whether or not the Duly River 
turtles represented merely uw non-breeding out- 
lier Of a parent Papuun population. AL speci- 
mens were relatively small, ranging in curapace 
length from about 26 to 38 erm, It Was also 
delermined from faeces that food ingested by 
one of them comprised figs and freshwater 
snails. 

A seeond verified record now cones from 
the South Alligator River system. approxi- 
mately 400 km eust of the Duly River site. 
There, in’ Yellow Waters billabong on Jim 
Jim Creck, a single female (CSIRO R. No, 
320) was ecuught by uw CSIRO fauna survey 
team on 35 November 1971. The specimen 
(Fig. 1). deposited in the museum of CSIRO's 
Division of Wildlife Research, Cunberra, is 
large, having a carapace Jength of 45.6 cm. 
This compares with ca 48-50.5 cm for the 
Jargest New Guinea specimens (Walther 1922: 
Schultze-Westrum) 1963). Other dimensions, 
taken from life, are: carapace breadth (includ- 


ing marginals) 36.5 cm: total height (carapace 
+ plastrog) 14.8 em: head length (to base of 
crown) 15 cm; head width 8.1 cm; heqd + 
neek length (to gular shields of plustron) 18.5 
en Tore-limb length (posterior margin of 
flipper) 26.4 em: hind-limbh length (anterior 
margin of flipper) 25.3 em: tail (to base of 
anal shields of plastrow) 16.9 em long. with 
12 or 13 dorsal scule bands. Soft part colours: 
shell. limbs. head and tail, mid to durk olive- 
brown dorsally, prading to cream ventrally 
(fleshy cream on plastron), iris mid blue-green- 
grey, Except for the reduced number of caudal 
scutes, quoted ut 14-16 for New Guinean 
specimens by de Roo (1915). the South Alfi- 
gator specimen appears to be identical with 
New Guinean forms. 

The condition of the reproductive tracts 
showed the female to be in the process af lay- 
ing: a large number of enlarged megalecithal 
ova Were present in the ovaries (Pig. 2). Many 
follicles also appeared ta have ruptured 
recently, and both oviduets were murkealy 
swollen. The large unshed Ova were jlmost the 
size of shelled eggs according to the dimensions 
for the latter illustrated by de Rooij (1915. fig. 
102). Because Yellow Waters billabong. c@ 30 
km up stream above tidal influence on the 
South Alligator system, had been landlocked 
between April and November during the mon- 
soonul dry season, there can be no doubt that 
egus had been deposited somewhere ulong the 
billabong, This represents the first evidence 
that the species breeds. in Australia. 

While kept alive for a lime in water. the 
turtle defaecated large quantities of partly 
digested husks of the stone Fruits of Purteinis, 
as Well ds a few shoot leaves (Melaleuied and 


* Division of Wildlife Research. CSIRO, P.O, Box 84, Lyneham, A,C.T., 2602, Ausiratia, 
! Known locally in the Northern Territory by the appropriate vernacular of “pig-nosed turtle” (J. 


Cin. pers. comm. }. 


Trios, R. Suc, S, Aust. Vol, 96, Part 2, 31 May 1972. 


116 R, SCHODDE, I. MASON, AND T. O. WOLFE 


1OcntT 


Fig. |. The Pitted-shelled Turtle. Caretiochelys insen/pta (CSIRO specimen R, No. 320) 


- 
7 . 
[een See Oe ee 7 = 


Fig. 2. Reproductive tract of ovulating Carettochelys insculpta (CSIRO specimen R. No. 320). 


PITTED-SHELLED TURTLE FROM AUSTRALIA W7 


Leguminosac spp,), seeds, roots. picces of 
aerenchymatous plunt slem, and traces of ani- 
Mal matter. ‘The animal matter, comprising ca 
| of the defaccated matertal, included fresh- 
Water snails (Thiaridae sp.), water-hoalmen 
(Corixidag sp.), the water beetles Hosheod view 
scutellaris Germ, (Dytiseidae) and A ydro- 
philus fatipalpus Cast. (Hydrophilidac). and 
ants (fridantyrmesx sp.). Upon dissection, the 
eolon and Juwer intestine of the turtle were 
found to be packed with Pandenus fruit husks. 
Pertunth segments remained attached to many. 
of the husks, indicating that the turtle had 
presumably -broken and euten hurd, green, 
fruiting cones with its juws—no mean feat. 
T G. Schultze-Westrum (pers. comma has 
also Observed the species feeding on pandan 
fruit in New Guinea. Thus, although turtles 
bave proved to be somewhat omnivorous 
{Schultze-Westrum 1963; Cogeer 1970; J. 
Cann, pers, comim.}, the fruits of Pandanus, 
whenever falling from trees of the Various 
species of the genus that commonly line and 
overhang the estuarine and lower freshwater 
reaches of rivers in both southern New Guinea 
und northern Australia, would appear to con- 
stitute a rather significant item of their diet. 
There have been numerous other records of 
“freshwater turtles’ in northern Australian 
rivers jn recent years (cf, Cogger 1970) and. 
though in-one case unsupported by specimens, 
at least two appear to be of authentic Carerro- 
chelvs insculpta. Qne is of specimens examined 
hy J. Cann (pers. comm.) from the upper 
teaches of the Daly River about 140 km above 
tidal influence in the Northern ‘Territory. A 
caripace of one of the specimens has been 


CReg. No. R.31717), The other, published inci 
dentally by St, John (1967, p, 527) and Parker 
(1971), is of observations made by the lute A. 
de Lestang who observed “herds of turtles” 
devouring the fallen fruits of Pundanus (P. de- 
lestangii St. John) in perennial rivers south- 
west of Burketown in north-western Queens- 
lund and/or eastern Northern ‘Territory. As 
all other freshwater chelonians in the region 
are curiivorous, it seems probable that de 
Lestung'’s observations cefer to Carerrochelvs. 

These records suggest that the Pitted-shelled 
Turtle occurs in the lower (to upper) reaches 
of major river systems across northern Austra- 
lia, at least from the Victoria River District af 
the Northern Territory to as far east us the 
Gulf country of Queensland and perhaps 
western Cape York Peninsula. From local in- 
formation. J. Cann (pers. comm.) believes the 
lure to Be rather common in most coastal 
rivers flowing into the Joseph Bonapatie and 
Van Diemen Gulfs, In averall distribution, 
then, the species apparently occuts in river 
systems along the landward margins of the 
Sahul Shelf, Little is yet known of its oecur- 
rence or movernents in the intervening Arafura 
Sea. 

Acknowledgements 

We are indehied to Messrs. W. Vestjens and 
I. H. Calaby, CSIRO Division of Wildlife 
Research, for assistance with measurements 
and examination of the CSIRO specimen and 
in the preparation of the text of this paper; and 
to Dr. T. G. Schultze-Westrum, and Mr. J. 
Cann of Yarra Road, Phillip Bay, Sydney for 
additional observations on Cerettachelys in- 
senipia. Mr. &. Slater, CSIRQ Division of 


placed in the Australian Museum, Sydocy Wildlife Research, took the photographs. 
References 
Coccrr, H. G. (1970)—First recard of the SeuunTze-Westaum, T. (1963).—Die Papua- 
Pitted-shelled Turtle, Caretrochelys prsculpta, schildkrote aus Neuguinea, Natur nad 


from Australia. Search 1, 41, 

De Koop), N. (1915)—"The reptiles of the tndo- 
Australian archipelago, 4, Lacertilia, Chelonia. 
Emydosauria.” (Brill; Leiden.) 

De Ron, N. (1922), -Reptiles (Lacertilia, 
Chelonia and Emydosauria), Neva Giined 13, 
4001... 133-153. 

Parnsik, S. A. (1971) —Association hetween the 
Sulphur-crested Cockatoo and Pandanus. 
WA. Naturalist 12, 23. 


Museunt 93, 119-127, 

St. Jonn, H. (1967)—Revision of the Genus 
Pandanus Stickman, Pt. 23, Three Australian 
Species of Pandanuyy, Paci{. Sei 21, 523-530. 

WALTHER, W. G, (1922)—Die Neu-Guinea- 
Schildkrote Carettachelys inseulpia Ranwsay. 
Nova Guinea 13, Zool,, 607-704. 

Wermurs, H., & Merrens, R. (1961 ),—“Schild- 
kroten, Krokodile, Bruckenechsen.” (Gustav 
Fischer: Jena.) 


THE MORPHOLOGY AND RELATIONSHIPS OF MUELLERENA 
WATTSII (HARVEY) SCHMITZ (CERAMIACEAE: RHODOPHYTA) 


BY ELISE M. WOLLASTON 


Summary 


The morphology and life history of Muellerenu wattsii (Harvey) Schmitz is described and its 
relationships discussed. It is recognized as belonging to the tribe Crouanieae (Ceramiales, 
Rhodophyta) on the basis of thallus morphology and stages in development of the procarp and 
carposporophyte. Features including regularity of branching pattern and elaboration in development 
of the involucre surrounding the carposporophyte suggest a probably phylogenetically advanced 
condition. 


THE MORPHOLOGY AND RELATIONSHIPS OF MUELLERENA WATTSII 
(HARVEY) SCHMITZ (CERAMIACEAE; RHODOPHYTA) 


by Fitse M, Woitaston* 


Summary 


The morphology and life history of Muellerena wattsit (Harvey) Schmitz is described and jts 
relationships discussed, It is recognized asx belonging to the tribe Crouanieae (Ceramiales, Rhodo- 
Phyta) on the basis of thallus morphology and stages in development of the procarp and carposporo- 
phyte, Featares including regularity of branching pattern and elaboration in development ot the 
involucre surrounding the carposporophyte sugeesr a probably phylagenctically advanced condition, 


Introduction 


Magllerena waist CAHarvey) ‘Sehmitz in 
Schmitz & Hauptfleisch was described as 
Crowenia watsii by Harvey (1863), who con- 
sidered it closely allied to C'. agardhiana [now 
Ptiloecladia upardhiana (Harvey) Wollaston 
1968). Schmitz (TASD) listed the species as 
helonging to a new genus Muellerella which, 
however, he formally described (in Schmitz & 
Hauptfieisch 1897) as Muellerena with M. 
witht as the type species. 


Muellerend is a monotypic genus scemingly 
quite distinct from other closely related genera. 
Schmitz (1889) placed it in the Dasyphileue 
but Kylin (1956) suggested that tt was pro- 
hubly more closcly related to Cronania and 
considered it to be insufficiently known for 
correct placement. Hommersand (1963), after 
cxamining specimens of M. warnsii in TCD, 
concluded that it was probably correctly placed 
in Dasyphileae, Elowever, detailed study of 
both veyetitive and reproductive features show 
thal it is best placed io the Crouanicae, 

Material used for investigation has been 
muinty drift plants collected at Stinky Bay, 
Nora Creinu, §. Aust. (Wollaston, 14.xi.1955; 
ADU, A20004: Wollusten, 19.v,1964; ADU, 
A27924) and at Seal Bay, Kangaroo T., S. 
Aust. (Womersley, 211.1965: ADU, A2ZB819), 
These collections inchided both tetrasporangial 
and carposporangial plants. 


Muellerena wattsii (Harvey) Schmitz in 
Schovitz & Hauptfleisch 1897: 496, De 
Tont 1903: 1388; $924; 490, Lucas 1909; 
50. Lueas & Perrin 1947; 344, Mazza 
1911: Na, 397. 


Crouania wartsii Harvey 1863; synop. No. 
637, pl, 291, J. Agardh 1876: 86, Tisdall 
T&98: 503. 

Muellerelle owatisii Schmitz 1889; 
(nomen nudum), Kylin 1956: 397. 


Challus to 13 cm high with terete, sparingly- 
branched axes beuring alternate. distichous 
lateral branches up to scveral em long und 
usually pinnately branched in the outer part 
(Fig. 1)% laterals borne from allernate axial 
cells, occasionally with # shorter branch (less 
thar | cm long) opposite or between the 
longer laterals (Fig. 2}. These shorter branches 
are initivted on the basal cell of an omginal 
whorl-branchlet and develop in its place. 
Axial cells aro 1-LS times as long us broad 
with cells of the central mature thallus usually 
350-400 ym long, Each axial cel) bears trom 
its upper part a whorl of 5 whorl-branchlets 
(Figs. aii, 4), with the exception that those 
cells which bear lateral branches often produce 
only 3 whorl-branchiets and | lateral branch 
(Fig. 31, ini). 

Growth takes place by transverse divisions 
of an apical cell and whorl-branchlets are ini- 
tiated usually on the sub-apical cell (Fig. 5) 
with the first-formed initial of each whorl in 
lateral position, the second and third to the 
right and Icft of it respectively and those last- 
formed oppasile the first one (Fig. 3ii).. During 
early development, lateral branches are charac- 
teristically curved due to their having the first- 
formed and hence the longest whorl-branchlets 
borne on the abaxial (outer) side, while the 
shortest most immature ones are adaxial in 
posilion (Fig, 5), However, after initial elonga- 
tion of 1—-several mm, each young lateral pro- 
duces at its tip further alternate. distichous 


451 


? Deparimenit of Botany, University of Adelaide. &. Aust. 3001. 


Trans, R. Soc. S, Aust. Wot. 96, Part 2, 31 May 1972. 


(20 ELISE M. WOLLASTON 


1 2 


Plant habit (Stinky Bay, Nora Creina, S. Aust. drift, Wollaston, 19.v.1964. ADU A27924), 
Type specimen, TCD (Warrnambool, Vic., cast ashore, Marts, Sept. 1860, Harvey 221), 


Regular. distichous arrangement of short lateral branches and occasional shorter branches 


4 
*\ “ 
Fig. 1, 
Fig, 2. 
opposite or between the longer ones. 
lateral branches on alternate axial cells, On 


each of these axial cells the lateral branch is 
initiated first followed by whorl-branchlets to 
the right and left of it and the last formed one 
opposite the lateral branch (Pig. 3i, ili, iv). 
Most lateral branches cease growth early bul ou 
few continue to elongate and form indeter- 
minate thallus branches (Fig. 2). Young cells 
enlurge rapidly and gland cells and. tetras- 
porangia may occur very close to branch tips, 


Mature whorl-branchlets consist of several 
consecutive die or tri-chotomous whorls of 
cells, and terminate in short, 2- or 3-celled 
chains of small cells, cach up to 7 yam diam, 
often terminated by a slender, elongate hair to 
[80 pm long (Fig. 4). Cells of whorl-branch- 
Jets ure 1-2 times as long as broad and up to 
60 pm Jong in the central part of mature 
whorl-brunchlets. Whorl-branchlets are com- 
monly lost trom older axes and particularly 
from the upper and lower axial face between 
the distichously-arranged branches, 


Axes ure corticated, except when very 
young, by descending, branched filamentous 
rhizoids of elongate cells which arise from the 
basal cells of whorl-brunchlets (Fig. 6) and 
intertwine to form a dense anial covering with 
short, horizontal branches, composed of a 
chain of several small cells, projecting out- 
wardly. In older parts of the thallus the axl 
cells become very thin-walled aod may be 
almost indistinguishable within the cortical 
cylinder. 

Ovoid to pyriform ghind cells up to 16 am 
long, evch within a thick gelatinous sheath, 
occur as homogencous, refringent structures 
horne in place Of outer branches of whorl- 
branchlets and scattered, sometimes bun 
dantly, over the thallus (Fig. 7). Cells of the 
thallus appear to be uninucleate although pro- 
perly fixed material has not been available for 
study with specific nuclear stains. Rhodoplasts 
vary from small and rounded in young cells lo 
reticulate and finally to elongute in mature cells 
(Fig. Si-iii), 


MORPHOLOGY AND RELATIONSHIPS OF MUELLERENA 21 


Procarp and Carposporaphyte—Carpogonial 
branches, 4-celled when mature, are initiated 
sinaly on uw supporting cell which ts one of a 
whorl of 4 cells borne from the upper part of 
the terminal cell of a short 2 (-3)-celled spectul 
fertile branch (Fig, 9). The fertite branch is 
produced at the outer end of the basal (or 
stcond) cell of a whorl-branchlet and repluces 
one branch of the normal di- or trichotomy. 
The sub-apical cell of the fertile brunch also 
bears a whorl of (4-)5 cells (Fig. 9). Bach 
fertile branch is initiated near the tip of a 
branch axis, so that a suceession of maturing 
procarps and carposporophytcs is produced as 
the branch axis elongates, Celle of the carpo- 
gonial branch are formed by transverse divi- 
sions of an initial cell which is cut off out- 
wardly from the supponing cell (Figs. 10-13). 
The lower three cells stain densely and appear 
homogeneous in structure while the carpo- 
gonium is smaller, often with a densely-staining 
protoplast concentrated im one portion of the 
eell, and bears an elongate trichogyne. Io 
90 pm long, usually swollen at its base and its 
tip (Figs, 12-14). A sterile cell is formed on 
the upper side of the supporting cell after ini- 
tiution of the curpogonial branch and is usually 
well-developed by the time the carpogonial 
branch is mature (Figs, Fl-13). 


Following fertilization. the carpogonium en- 
larges jind becomes rounded in form while the 
uichogyne degenerates and an auailiary cell 
develups from the upper part of the supporting 
cell (Fig. 15). At this stage the three stevile 
cells, making up the whorl which includes the 
supporting cell of the carpogonial branch, 
conimence to enliurge und ench becomes 
roughly triangular in shape; the sterilo cell 
berne on the supporting cell divides to form a 
chain of several cells (Fig. (5) while the cells 
forming the whorl ov the sub-apicel wxial cell 
elongate and produce terminally the first cells 
of branched involucral filaments (Fig. 16), 
Fusion takes place by means of a connecting 
cell between the carpogonium wand the upper 
part of the auxiliary cell, leaving only one or 
two small cell fragments in place of the carpo- 
gonium on the degeneraling carpogonial branch 
(Fig. 17}. 

Branched involuctal filaments formed. from 
the enlarged sterile cells on the apical and sub- 
apical uxial cells of the fertile branch develop 
rapidly, and loosely surround the developing 
cntposporophyte (Fig. 17). The hhasal cells of 
the upper whori of flaments remain charac- 
teristically triangular and larger than other cells 


of these branches (Fig, 17). The auxiliary cell 
cuts off a gonimoblast cell fromi its upper side 
and simultaneously forms 2 pil-connection with 
the apical cell of the fertile branch axis (Fig. 
18). Through this connection mutriment ix. pos- 
stbly conveyed more directly to the carpo- 
sporophyte, while the old supporting cell acts 
as the basal cell of an involucral filameni. 
Gonimolobe initials, which each give rise to a 
rounded group of curposporangia, develop suc- 
cessively with the first one or two gonimalches 
produced in a lateral position, Further goni- 
molobes are produced without regular order 
30 that a total of 6 of more groups of cgarpo- 
sporangi wt various stages of development 
may be present at the one time (Fig. 19), As 
the first carposporangia mature, the newly 
formed pit-connection between the axial cell 
and auxiliary cell gradually widens and the 
connection between the lower part of the aus 
liary cell and the supporting cell remains small 
and peobably non-functional or is finally 
broken (Fig. 19), The involucral filaments, 
each branched several times, curve upward and 
loosely surround the mature carposporophyte. 


Spermarangia—not recorded, 


Tetrasporangia— Sphencal,  tetrahedrally-di- 
vided tetrasporangia, seldom greater than 25 
pm diam, are home on the outer cells of 
whorl-branchlets in place of vegetative 
branches {Fig, 20), in a similar position to 
glund cells. They may occur of aity pal't of the 
thallus but are usually most abundant on young 
brunches. 

Type Locality—Warrnambool, Vic. ( Wars, 

Sep: 1860). 

Holotype—TCD, Harvey Alg. Aust. Exs. 

No. 221. 

Disiribeion—From West I, and Kangaroo 

I,. 8. Aust.. to Warrnambool, Vic. 


Diseyssion 
Muellérene wellsit is characterized hy the 
following vegetative and reproductive features: 

(a) a consistent branching pattern and short 

Iwterul branghes developed regularly from 

the outer end of axes, 

whorl-branchlets in whorls of 5 on each 

axial cell, except on those which bear 

short laterul branches, 

(c) fusion between the lower part of the auxi- 
liary cell and the fertile axial cell during 
carposporophyte development, 

(d) a distinct filamentous involuere surround: 
ing the carposporophyte and involying the 


th) 


122 ELISE M. WOLLASTON 


original supportilig cell of the catpogonial 
branch, 

Vegetative features such as the alternate-dis- 
ichous srrangement of shurt Literal branches 
at the tip of each prevyiously-formed lateral, 
the form of axial corti¢alion, und the consis- 
tency in arrangement and number of whorl- 
branchlets on @uch uxial cell, suggest a telu- 
tionship with the Prifocladia group of the 
Ciousniexe. These features were tegarded by 
Wollaston (1968, p. 404) as indicative of a 
phylogenetically advanced thallus form. Hom- 
mersand (1963) stated that Muellereng wattsii 
was quadriverticillute und he considered the 
order of initiation of whorl-branchlets in a 
thodomeluceyn sequence (the first abaxial, the 
next two to the right and left of the first and 
the fourth one adaxial) to be a significant taxo- 
nomic feature characteristic of the Dasy- 
phileae. However, M. waftsii has in fact 5 
Whorl-branchlets per whorl except where short 
Jateral branches are produced, and in these 
Whorls, the order of development of the 4 


ifitixls could have adsen from a crowanioid 
seyuence in which the second branchlet is 
formed opposite the first and the third and 
fourth at right angles lo them, Suppression of 
the adaxial branchlet when adjacent to another 
axis is commonly tound in a number of taxa, 
for example, in species of Plaryrhanuion J. 
Agandh and Amoenothamnion Wollaston 
(Wollaston 1968). In Muellerena waersit this 
could explain development of the intermediate 
whorl-branchlets of cach whorl prior to initia- 
tion of the adaxial whorl-braunchler{s), which 
al fimes is completely lacking at the base of 
lateral branches. M4. wenesii clearly evolved a 
stable pattern of branching and on this basis is 
probably vegetatively advanced. 

Several other vegetative features of M. 
warrsil also suggest telationship with the 
Crouaniewe group. Gland cells, not previously 
recorded for A¢_ wartwsii, aré similar in form to 
those found in Prilucladia australis (Plarv.) 
Wollaston, P. vestita (Hary.) Wollaston, and 
Gulvonia annulate Harvey, although they lack 


position of fusion with connecting cell from carpo- 
1 place of carpogonium on degenerating carpogonial 


Tig. 3. tiv. Arrangement and sequence of initiation of short lateral branches (Lj and whorl-branch- 
lets in whorls on successive axial cells near the tips of axes. (Diagrammatic.) i, ui, iii repre- 
sent in transverse section the 2 celis shown in iv. 

Fig 4, ‘Vransverse section of axial cel! bearing a whorl of 5 whorl-branchlets with tetrasporangia. 

Fig. 5, Tip of branch sxis showing altcrnale, distichous arrangement vf youne lateral branches 
(Ly-Ly) on alternate axial cells and abaxial initiation of first-forroed whor)-branchlets on 
cells of Jateral branch ates. (Whotl-branchicts on faces of axcs omitted for clarity.) 

Fig. # Benanched, descending cortical rhizoids borne on basal cell (6) of a whorl-branchlet. 

Fig. 7, Gland-cells borne im place of branches of whorl-branchlet. 

Fig. 8 Rboduplast structure (i) young cell with rounded chodoplasts, (it) enlarging cell with reti- 
culate rhodoplasts, (iii) mature cell with elongate rhadoplasts. (Diagrammatic. ) 

Tig. 9. Carpogonial branch on special 2-celled fertile branch borne in place of a whorl-branchict 
brinch on basal (or second) cell of whorl branchlet. 

Fig, LO. Carpoyonial branch initial eut off ounyardly fram supporting cell, 

Fig. |]. Young curpogunial branch, 3-celled stage, on supporting cell Which also bears a small 
aterile cell (5), 

Fig. 12. Curpogonial branch, 4-celled, with developing trochogysuc. 

Fig. 13. Carpogonial branch with fully elongated trichogyne. 

Fig, 14, Fusion of spermatium (sp) with mature ctrichogyne 

Fig. 15. Auxiliary cell (4) formed from upper side of supporting cell; curpogonial branch with 
enlarging carpogonium (c) and terminal remnant of trichogyne; sterile cell (s) bearing first 
cells of involucral filament. 

Fig. lé. Carpogonium (c) enlarged just prior to fusion with auxiliary cell (a); involucrsl filaments 
commencing to form from cells, including the supporting cell, of whorls an axial cells (axy, 
ake) of the fertile branch, 

Fig, 17. Pratrusion on auailiary cell (a) markin 
gonium; small cell fragment remaining 
braach: miurked increase. in develupinent of involucral filaments. 

Fig. (2. Formation of pitconnection between lower part of auxiliary cell (a) and axial cell (ax) 
prior to breaking of connection between quxiliary cell and supporting ecll; initials of 2 lateral 
groups of carposporangia formed on gonimoblast cell (2). 

Fig. (9. Enlarged fusion between lower part of auxiliary cell (a) and axial cell (ax,); succession of 
carposporangial groups forming on gonimoblast cell; supporting cell free from carposparo- 
phyte. bearing involucral filament. 

Fig- 20. Tetrasporangia and gland-cell borne in place of vegetative branches of whorl-branchlet 


MORPHOLOGY AND RELATIONSHIPS OF MUELLERENA 


“o 
mm 
ul 
nS 
~ 
SS 
& 
ur 
ou 
a 
E 
a 
2 


3 
2 
2 
a 
re 
! 
2 
wo 
g 
= 
2 
2 


124 


ihe crystallike inclusions recorded for these 
species. Branched cortical filaments bearing 
short ollwardly-orientated chains of cells are 
similar to thase found in Prilectadia palchra 
Sounder and a tendency towards distichous 
branching nf the thallus, well defied in 

wellerend watisi, is also characteristic of the 
Crouanieae group and is best developed in 
species considered to be phylogenetically ad- 
vanced, 

Development of the procanp and carpo- 
speraphyte also biusically resembles that found 
in gerera of Crouanieae. The 4-celled carpo- 
gontal branch is borne on a special fertile 
branch as in Gilsonta. The connecting cell in- 
volved in fusion between the ¢arpogonium and 
auxiliary cell is much larger in Muellerena 
watesii than in genera of Crouanieue, but yub- 
sequent development of the carposporophyte 
with futeral initiation of the two first-formed 
groups of carposporungia is similar ito that 
seen in species of Plilecladia, Gulsonia and 
Euptilocladin Wollistan. Mitellerener wattsii 
differs, however, in elaboration of the involucre 
which surrounds the carpasporophyte and in 
the secondary development of a pit-connection 
linking the lower part of the auxiliary cell ro 
the axial cell upon which the procarp was 
developed, Following this fusion, the original 
connection between the supporting cell and the 
lower part of the auxiliary cell is ustially 
broken so that the supporting cell functions as 
an eilarged basal cell of an involucral filament 
similir lo its sister-cells of the whorl. The in- 
yolucral lament bome on the supporting ceil 
ts Inithved as a sterile cell on the supporting 
cell during enlargement of the carpogonial 
brunch amd elongates at whout the same dime 
as the other involucral branches commence to 


ELISE M. WOLLASTON 


develop. These events probably allow a better 
nutritional supply to the carposporophyte while 
at the same time providing for development of 
the filamentous involuere. Although Mfiel- 
lerena watts differs from) species of Prilecladia 
in having a more consistent branching pattern, 
§ whorl-branehlets pee whorl, fusion between 
the auxiliary cell and fertile axial cell and a 
more claborare involucre surrounding the 
carpasporophyte, the two genera mre basically 
similar in both vegetative and reproductive 
features, This similanty was noted by De Toni 
(1903) when he placed two specics now recog- 
nixed as Prifocladia pulchra Sonder and P. 
agardhiana (Harvey) Woll. in the genus Muel- 
lerena. M, watisit is also samilar lo Gulsonia 
in the presence of gland cells and the develop- 
ment of 4 special Fertile branch bearing the 
procarp and, later, the carposporophyte. Thus 
it seems likely that Priloctadin, Gulsonia and 
Muellerena are closely related and MreJleresa, 
showing greater consistency in vegetutive feu- 
tures. and elaboration in carposporophyle 
organization, is phylogenetically the most 
highly advanced. The singe of Features already 
known for gencra of the Crouanieac covers 3 
possible evolutionary sequerice leading to the 
increased organization and stability of thallus 
features characteristic of Muellerena. No simi- 
lar relationship can be traced in the Dasy- 
phileae or other group of the Cerannacese and 
it thus seems logical to include Muellerena in 
the tribe Crouanieae of the Ceramiacewe. 


Acknowledgements 
Tam grateful for the loan of specimens Crom 
the National Herbarium, Victoria, and for 
technical assistance provided by a grimt from 
the Australian Research Grants Committee. 


References 


Acarpu, J, G, 19876),—Species, Genera et 
Ordines Algarum 3 01), pp. 1-724. Eperisis 
Systemulis Plorideurum. (Lund.) 

De Tost, J. B, (1903) —Sylloge Algarom omaiim 
hucusygue Cognitarum 4. Florideac, Sect. 3, 

p. 775-1521. (Padua, ) 

De ‘emt, J.B. (1924).—Sylloge Alvarum omnium 
hucusque Cognitarum 6. Floridese. (Padua) 

Harvey, W. H, (1863) —Phycologla Australica 5, 
Plates 241-300, synap. pp. [-73. (London.} 

Hom™erasann, M, H. (1963).—The marpholosy 
ant classification of some Ceramiuceac and 
Rhodontelaceae. Univ. Catfif. Publ. Bot. 35 
(2), 165-366. 

Kris, H. (1956)—Die Gattungen der Rhode 
phycesn, (Lund.) 

Lucas, A. H. S&S (1999).—Revised list of the 
Foucoideae ond Florideac of Australian. Proc, 
Litin, Sac. NSM. 34, 9-40. 


Lucas, A. HH. S.. & Prerin, F, (1947).—The Sea- 
weeds of South Australia. Part I The Red 
Scaweeds, pp 107-458. (Govi. Printer: Ade- 
laide.} . . 

Mazza, <A. (191b)—Sageio di Algologia 
Oceanica, Nueva Novairiste 22, Nas, 369-414, 

Scumirz, F. (1889)--Systematische Uebersicht 
der bisher bekannten Gattungen der Flori- 
deen, Flore 72, 435-456, pl, 21, 

Sonmirg, F., & Hauereveiscu, P. (1897 ).—Cera- 
miacene. 7 A. PNczer & K. Paanti, "Die 
Natirlichen Pflanzenfamilien” I (2), 481- 
504. (Leipeig.) 

Tisoaty, 1. T. (1898)—The algae of Victoria. 
Rep, 7th Meet, Aust. Ass. Adv. Sci,, Sydney, 
1898, pp. 493-S)6. 

Wottaston, E. M. (1968)—Morphoiogy and 
taxonomy of southern Australian geners of 
Crouanieae Schmitz (Ceramiaceac, Rhode 
phyta). Aust, J. Bot. 16, 217-417, pts. 1-10, 


VOL. 96, PART 3 31 AUGUST, 1972 


TRANSACTIONS OF THE 


ROYAL SOCIETY 
OF SOUTH AUSTRALIA 


INCORPORATED 


CONTENTS 


Smith, Meredith J. Small fossil vertebrates from Victoria Cave, Naracoorte, 
South Australia. Il. Peramelidae, Thylacinidae and Dasyuridae 
(Marsupialia) - - - - - - - - - 125 


Mawson, Patricia M. The genus Acuaria Bremser (Nematoda: Spirurida) in 
Australia - - - - - - - - - - 139 


Milnes, A. R., & Bourman, R. P. A Late Palaeozoic glaciated granite surface at 
Port Elliot, South Australia - - - = = = - 149 


Tyler, M. J., & Parker, F. Additions to the hylid frog fauna of New Guinea, with 


description of a new species, Litoria timida - - - - 157 


a 


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


SMALL FOSSIL VERTEBRATES FROM VICTORIA CAVE, 
NARACOORTE, SOUTH AUSTRALIA 


Il. PEAMELIDAE, THYLACINIDAE AND DASYURIDAE (MARSUPIALIA) 


BY MEREDITH J. SMITH 


Summary 


Abundant fossil remains of marsupials and rodents have been found in Victoria Cave, near 
Naracoorte, South Australia. The presence of certain large, extinct herbivores in the assemblage 
suggests that the deposit may be of Pleistocene age. This paper describes remains of Jsoodon 
obesulus (Shaw, 1797), Perameles gunnii Gray, 1838 and P. bougainville Quoy & Gaimard, 1824 
(Peramelidae); Thylacinus cynocephalus (Harris, 1808) (Thylacinidae); Dasyurus maculatus (Kerr, 
1792), D. viverrinus (Shaw, 1800), Antechinus flavipes (Waterhouse, 1838), A. swainsonii 
(Waterhouse, 1840), A. stuartii Macleay, 1841, Sminthopsis murina (Waterhouse, 1838) and S. 
crassicaudata (Gould, 1844) (Dasyuridae). Extensions of the previously known ranges of P. 
bougainville and A. stuartii are noted. 

The larger species are represented mainly by juveniles and it is suggested that the small mammal 
remains were accumulated by owls. 


SMALI, FOSSIL VERTEBRATES FROM VICTORIA CAVE, NARACOORTE, 
SOUTH AUSTRALIA 


Il, PERAMELIDAE, THYLACINIDAE AND DASYURIDAE (MARSUPIALIA) 


by Merepity J. SMita* 


Summary 


Abundant fossil remains of marsupials and rodents have been found in Victoria Cave, near 
Naracoorte, South Ausiralia. The presence of certain Jarge, exlinct herbivores in the assemblage 
suggests that the deposit may be of Pleistocene age. This paper describes remains of Isoodon obesulus 
(Shaw, 1797), Perameles gunnii Gray, 1838 and P. bongainrville Quoy & Gaimard, 1824 (Perameli- 
due); Thylucines eynocephalns (Harris, 1808) (Thylacinidae); Dasyurus maculatus (Kerr, 1792), D, 
viverrinus (Shaw, 1800), Aatechinuy flavipes, (Waterhouse, 1838), A. swainsonii (Waterhouse, 1840), 
A.. stuartii Macleay, 1841, Sminthopsis muring (Waterhouse, 1838) and S. crassicuudata (Gould, 1844) 
(Dasyurjdac), Extensions of the previously known ranges of P. howgainville and A, stuartii are noted, 
The larger species are represented mainly by juveniles and it is suggested that the small mammal 


remains were accumulated by owls. 


Introduction 

Victoria Cave, m Tertiary limestone near 
Naracoorte (lat, 37°0°S, long. 149°48’E) has 
been Open to tourists for many years, In 1969, 
the Cave Exploration Group of South Aus- 
tralia (CEGSA) discovered further extensive 
rumifications of the cave and. in one chamber, 
a silt deposit containing abundant skeletal re- 
mains of large animals. Many of these were 
later identified as remains of extinct marsupial 
herbivores (sthenurines and diprotodontids) 
and of the marsupial lion, Thylaceleo sp. 
(Wells, pers. comm.). The sthenurines and dip- 
Totodontids are believed to have become extinct 
at the end of the Pleistocene (Tedford 1967), 
and the deposit in Victoria Cave is therefore 
probably of Pleistocene age. 

Bone chips occur in cores taken as decp as 
2.5 m but the maximum depth of excavation 
at present is 80 cm. 

The potoroincs (Macropodidaé), petaurids 
and burramyids have been described previously 
(Smith 1971); the present paper describes the 
peramelids, a thylacinid and the dasyurids. 


Methods 
The methods of sieving the bony remains 
from the silt, and their subsequent cleaning 
and preservation have been described (Stith 
1971). Measurements of teeth have been made 


in the way described in that paper, with the 
exception that, in the peramelids only, the 
maximum. anteroposterior Jengths of mandibo- 
lar molar teeth were measured on the lingual 
side. (The slope of the anterior cingulum from 
lingual side to buceal side hindered accurate 
Measuring on the buccal side.) Additional 
mandible measurements. were taken as follows; 
Length of ascending ramus: Distance between 
anterior and posterior borders of the ascending 
ramus, from the midpoint of the posterior 
border and perpendicular to the ramus mid- 
line. 
Breadth at M%,; Thickness of mandible below 
Ms. 
Height at M4; Distance from alveolar margin 
at middle of M4, to inferior border of manuible, 
and perpendicular to the infcrior border. 
‘The taxonomy used is that of Ride (1970) 
unless stated otherwise. 


Family PERAMELIDAE 
Tsoodon obesuliis (Shaw, 1797) 

The following features were used to dis- 
tinguish fragmentary remains of [soodon from 
Perameles, 

(i) The hypocone of each maxillary 
molar (except M4) is well developed 
in Tseadon so that in horizontal sec- 
tien these teeth appear as rounded 


*48 Leahreok Drive, Rostrevar, §. Aust. 5073, 


Trans. R. Sac. S. Aust. VoL 96, Part 3,31 August 1972. 


126 


blocks. In Perarieles the hypocones 
are much smaller and M}, M2, and 
MS appear as truncated triangles, 
tapering lingually. The molar alveoli 
refiect the shape of the {ccth. the 
lingual root relative to the buccal 
length of the tooth being much longer 
in Jsoodon than in Perameles. 
In the mandible of Iseodon, the an- 
terior cdge of the ascending ramus 
makes an obtuse angle with the hori- 
zontal ramus, whereas ii Perameles 
the horizontal and ascending portions 
of the ramus join in a continuous 
smooth curve (Merrilecs 1965) (Figs. 
1. 2, 3). 
In Iseodon the lingual extremity of 
the anterior cingulum is almost as 
high as the apex of the paracanid of 
Mi, M4, and M4, whereas in Pera- 
meles the greatest height of the an- 
terior cingulum is much less than that 
of the paraconid, 

Tscodon was not abundant in the deposit and 
adults and juveniles were about equally rep- 
resented (Table 1), 


(ii) 


(iit) 


MEREDITH J. SMITH 


Modern specitnens of £ vhesulus in the 
South Australian Museum vary greatly in size, 
and wide variations occut even in adult speci- 
mens of the same sex and locality. The length 
of M!—3 varied from 9.2 to 11.4.mm (mean 
10.19, sd. 0.72) in eight South Australian 
inainland specimens and the length of M4—4 
from 12,7 to 15.8 mm (mean 14.49, s.d. 1.08). 
Victoria Cave specimens are smaller than 
modern mainland speciinens (Table 2) and 
are almost as. small as the insular subspecies, 
I, 0, nautieus where, in nine specimens, the 
length of M14—% ranged from 8.4 to 9.5 mm 
(mean 8.68, s.d. 0.35) and the length of M4—4 
from 11.8 to 12.5 mm (mean 12.20, s.d. 0.28). 
The teeth of Victorin Cave Jsooden are mor- 
phologically similar to those of modern /. 
ohesulus, 

Wakefield (1966b and in Mulvaney et al. 
1964) referred to a distinct, small form of 1 
obesulus from Mildura and from the Fromm’s 
Landing archaeological excavation on the River 
Murray, but he gave no measurements of this 
form, nor of the “much larger form abundant 
in S. Victoria’. Pleistocene specimens of J. 
obesulus fram Mammoth Cave, Western -Aus- 


TABLE | 


Mandibular and maxillary fragments of peramelid species found in the Victaria Cave, Naracoorte. Many 
isolated teeth were collected but have not heen ineluded in the table. 


Maxillae Mandibles 
Species Adult Juvenile* ~ Adult Tuvenile* Minimum no. 
Right Left Right Left Right Left Right Left of individuals 
Isoodon obesulus 4 4 os — 8 10 7 4 17 
Perameles gunnii 9 7 1 — 16 23 a4 46 77 
P. bouguinville Il 7 5 8 30 38 32 27 70 
Not determinable 2 2 3 3 i 6 9 5 15 


*A specimen was considered to be juvenile if P4 and/or M4 were not fully erupted. 


TABLE 2 
Some dimensions of mandibles and teeth of Isoadon ohesulus fram Victoria Cave. 
Number of Coefficient of 
Dimension Specimens Range (mm) Mean (mm) Standard error variation 

Length of ascending 

Tamus 7 6.3—F.3: 6.79 0.146 5.7 
Breadth at M& 6 242.8 2,58 0.075 TA 
Height af. Mé 6 3,6—4,5 4.10 0,152 9.1 
M}-4 iength 3 9.0—9.2 9.10 0.057 11 
P4 length 5 2.3—2.6 2.44 0.060 3.5 
P4 breadth 5 1.2—1.5 1.38 0.049 19 
M4-4 length 6 11.7—12.8 12.27 0,158 32 


SMALL FOSSIL VERTEBRALES FROM NARACOORTE II 12 


jratia, sre similar in size to modern specimens 
from the same area (Merriiees 1965) 

/, obesulus still occurs in the Naracoorte 
district, 

Genus PERAMELES 

Perameles was reptcscoted by many toothless 
mandibles and a few looth-bearing maxillae 
and mandibles. Adult mandibles, in which P4 
and M4 were erupted, could clearly be separa- 
ted on size into two species, distinguished by 
depth and thickness of the mandible, length of 
ascending ramus, und length of teeth (compare 
Figs 2 and 3; and Tables 4 and 5), Juvenile 
‘mandibles of the two. species overlapped in size 
but could be separated by the Jenpgth, und 
especially by the width, of their molar alveoli, 
Maxillae, hoth adult and juvenile, were identi- 
fied by the size of the molar alveolt. 


Perameles gunnii Gray, 1838 

Live specimens of the two large long-nosed 
bandicoots, P. granii and P. nasute Geoffroy, 
1804, appear quite dissimilar, the rump of 
guna being barred and that of nasufa being 
uniformly coloured (Ride 1970), However the 
skulls of the two species are similar in mor-~ 
phology. size and proportions, 1), provides the 
main difference, 1% of guntnii being double 
Tooted, antero-posteriorly long and bucco- 
lingually compressed, whereas 15 of rasrta is 
single rvoted and caniniform (Freedman 1967), 
This diagnostic foature could not be used on 


“4 


Victoria Cave material as no premaxillae were 
preserved, Additional differences are that the 
mandible is more slender in P. guanti, M4 is 
shorter buccally and M4 is shorter antero- 
posteriorly (Table 3). The figures given in 
Table 3 do not confirm Tate’s (1948) state- 
ment that P+ js nich broader in P. gunnii. 


Remains of the large Perameles from Vic- 
toria Cave conform in size with modem P. 
gunn {Table 4) and no morphological dif- 
ferences were detected between fossil and 
modera specimens. Remains of juvenile P- 
gurni? Were relatively abundant, but few adult 
specimens were found (‘Tahle 1), 


P. cunnii is not included in a list of the 
modern native mammals of South Australia 
(Aitken 1970) although three specimens of P. 
gunn in the South Australian Museum are 
registered as from South Australia (M1607 
from Mt. Gambier, M1613 from “‘Suuth Aus- 
tralia” and M3956 from the Rocks, Kougal, 
south-east of South Australia). All were collec- 
ted between 1891 and 1893. The present range 
of P. gunnié is southern Victoria and Tasmania 
(Ride 1970), 


Remains of P. gunnii were found in an 
aboriginal midden at Mt. Burr, South Australia 
(Finlayson 1966, unpublished!) and in a late 
Reeent deposit in the Bat Cave at Narucoorte 
(Tidemann 1967), 


TABLE 3) 
Some slienensions ef mandibles and teeth in: which Perameles nasuta differs fram P. gunmil. These 
lignres were celculated fram data given in Tables 2,3A and 3B of Freedman & Joffe 1967a and Tables 
1, 2A and 28 of Freedman & Joffe 1967b. 


Perametles nasuta 
95% confidence 


Dimension cxamined 


Perameyles punnii 


95% confidence 


N Mean (mm) limits of mean N Mean (mmj limits Of mean 

Length of ascending 

TaimMus 69 8.79 §.52—9.06 4) #34 4,23—6.46 
Breadth ut Mé 7 3.54 3.43—3,65 41 7.81 2.76—21.90 
Height at Mi 71 6.71 6.49—f.91 41 S88 §.70—6.06 
PS length 65 375 3.65—3.$4 40 3.44 3.9A7—2.45 
PS width hi 2,10 2,06—2,15 40 413 2,09—1,17 
MA width 81 271 2.6f—2.75 42 2.75 2,68—2,82 
M+ Buccal length 81 3.63 3.56—3.70 a3 3.09 3,053.15 
M¢# Lingual length 82 4,53 1.50—1.56 43 144 1.40—1.48 
M4 Anterior width o] 2,37 2.34—2.39 46 2.16 2.13—2.18 
M4 Posterior width R6 1.60 1,58—1.63 46 1.51 1.48—1,55 
M4 length 85 4.64 4.58—470 45 4.01 3 56—4,06 


‘Finlayson. H. H, i Campbell, T. D,, Edwards, R. & Hossfeld, P. S, (1966).—Archaeological excava- 
lions in the Southeast of South Australia. 24 pp. Transcript, Australian Instituie of Aboriginal Studies 
Library, Canberra, 


128 


Perameles bougainville Quoy & Gaimard, 1824 

The small bandicoots from eastern, central 
and Western Australia have been described as 
several different species, but Tate (1948) sug- 
vested that P. bougainville, fasciata, notina and 
eremiana might be local races of a single wide- 
spread species. Generally this has been accepted 
(e.g. Wakefield 1966a) although Ride (1970) 
retained eremiuna as a distinct species. Mean 
measurements of skulls and teeth are mostly 


MEREDITH T. SMITH 


larger in the south central population, noftina, 
than in the western population, bougainville 
(sens, strict.), but few of the differences are 
significunt (Freedman & Joffe 1967b). 
Remains of the small species of Perameles 
from Victoria Cave are similar in size and 
morphology to. modern specimens of P. bou- 
gainville in the South Australian Museum and 
to specimens from the Fromm’s Landing 
archaeological excavation (Table 5), 


TABLE 4 
Comparisons of somé dimensions of teeth and mandibles of Pérameles gunnii from Vieteria Cave, with 
those af a modesn sample fram Tasmania, CW. — Coefficient of variation. 


Modern P. gunnii from Tasmania 


P. guanii from Victoria Cave 


Dimension Standard Standard 
n Range(mm) Mean error CV. n Range (mm) Mean error C.¥. 

M+ width 10 3.6—4.1 3.91 .OS7 4.6 3 3.6—-3.7 3.63 33 1.6 
M+ buccal Jength 10 2,.8—3,2 3.03 032 3.3 3 293.0 2.97 033 1.9 
M+ lingual length 10 1.3—1,6 1.47 037 7.9 3 1.0—1.3 1.20 100 14.4 
M!—% alveolar 

length 10 1L4—-12.4 11.86 105 2.8 4 it.4—12.4 11.80 245 4.2 
Length of aseend- 

ing ramus 10 6.1—7.4 6.70 127 6.0 10 5.2—6,.2 5,75 100 5.5 
Breadth at M% 10 2.6—3.2 2.99 060 64 9 2.3—2.8 2.54 O55 6.6 
Height at M6 10 5.8—7.6 6,66 153 7.3 8 4.5—64 5.57 [90 9.7 
My alveolar 

length 10 3,.6—4,1 3.90 056 4.5 9 3.4—4.0 3.73 ATR: 7A) 
M4-4 alveolar 

length {0 14.6-—16.6 15.57 172 3.5 5 15.0—15.5 15.18 O86 1.3 

TABLE 5 


Comparisons of some dimensions of teeth and mandibles of Perameles bougainyille from Victoria Cave 
with these from Fromm’s Landing (specimens collected in levels 0-9). 


_ PF. hougainville from Fromm’s Landing 


P. hougainville from Victoria Cave 


Dimension Standard Standard 
n Range (mm) Mean error C.V_ on Runge fmm) Mean error C,V. 
M1-# Jenpth _ = oo — 4 91—10.4 9.40 334 70 
Longth of ascend- 
ing Tamus 8 3.8. 3.5 4.65 201 12.2 16 4.4—5.6 4,99 082 6.5 
Breadth at M& 18 2.0—2.7 2.24 044 84 35 1.8—2,5 2,16 043 10.6 
Height at Mi 16 3,4—5.4 4.46 128 115 24 3.6—5.4 4.33 15) «13.0 
P4 length 6 2.42.8 2.58 060 57 7 2,2—2.8 2.66 081 81 
Mj length 2 2.93.0) 2.95 50 24 5 2.73.2 2.96 O87 4.6 
M4 post. width 3 1.8—2.0 1.90 058 5.3 § 2.0—2.3 2,12 58 6,2 
Mé length 5 3.0—3.3 3.18 058 4.1 R 3,2—3.4 3.48 049 4.0 
M. post. width 4 2.1--2.4 2:25 065 5.7 8 2.22.4 2.34 032 3.9 
M3 length 8 3,0—3,4 3.19 058 5,2 8 3.0—3,4 3,23 041 3.6 
M*% post, width 8 19—2,2 2.09 .040 5.4 & 2.4—2.3 213 -037 4.9 
4 length 5 3.23.5 3,32 049 3.3012 3.2—4.0 3.47 067 6.7 
M4 post. width 5 .9—1,.4 1,20 N95 17.7 12 1.0—1.4 1.19 O31 91 
M44 length 3 11,9—12.4 12.1 152 2,2 S [2.4 —-13,7 12.88 218 3.8 


SMALL FOSSIL VERTEBRATES FROM NARACOORTE I 


OF twelve Victoria Cave eXamples of My. 
one showed a shor but distinct anterior cingu- 
lar shelf, six showed a slight Uepression in the 
anterior buccal region and tn fivé the unterior 
wall Was -contlnuously smooth Merrilees 
(1965) found that one Victorian specimen and 
two of fifteen from Western Australia showed 
the anterior-buccal depression on M4, On the 
basis of one Viclorign specimen of P. gummrii, 
Merrilees (1965) believed that a small cingular 
shelf on Mi might be characteristle of that 
species, but this is mot confirmed by my 
examination of eleven Tasmanian specimens in 
the South Australian Museum, in which sone 
has a distinct cingular shelf and only three have 
an saterior-buceal depression, 

P. bougeinville was equally abundant in the 
deposit as P. gwnnii, but a much higher pro- 
portion of FP. dowguinville, the smaller »pecies, 
wis adult (Table 1), 

The eastern Australian range of P, bougain- 
ville at the time of European settlement is 
poorly known. It occurred on the Liverpool 
Plaing in exsstern New South Wales (Ride 
1970) and it was apparently abundant near 
Mildura on the River Murray (Wakefield 
19460), its remains were found in an owl 
pellet accumulation of uncertain age in the 
Grampians, Victoria (Wakefield 1963), but not 
in other western Victorian cave deposits 
(Wakefield 1964), In the Fromm’'s Landing 
archaeological excavation, it was found from 
surface level w layers radiocarbon-dated at 
2105 = 8S years BP (Wakefield iz Mulvaney 
et al. 1964) but it was not represented in the 
Mt. Borer archaeological excavation, although 
Po curndl and J. ebesnlis occurred there (Fin- 
layson 1966 [footnote !] and personal observa- 
tions), The Victoria Cave specimens therefore 
extend ihe known range of the species in 
former times into south-costern South Austra- 
lia. 

Tn Western Australia, P. Aougatnville sur- 
vives today only on Bernier and Dorre Islands 
in Shark Bay (Ride 19707, However, its re- 
mains have been found in several caves on the 
Nullarbor Plain and along the southern half of 
the west coast of Western Australia (Lumdelius 
1960, 1963) and in a Pleistocene deposit tn 
Mammoth Cave (Merrilecs 1965). It occurred 
as a modem species in central and western 
Australia (Ride 1970). 


Family THYLACINIDAE 
Thylacinus cynoceplalus (Harris, 1808) 


Three isolated teeth are tentatively assigned 
to this species. 


129 


S.A.M. P)6120d is a worn canine, probably 
from a right mandible, with much of the 
enamel broken away and the root broken, The 
alze (max. anterd-pusterior length of root 11.3 
mm. max, width of root $5 mm) is similar 
to that of modern specimens of T.. cyrocepha- 
tus an the South Australian Museum, In the 
modern specimens the crown is not severely 
worn, for the upper and lower canines do not 
meet directly; but the crown of the Fossil tooth 
has either been severely worn, or was broken 
befure death, There is no anterior wear surface 
such as ovcurs in many modern specimens of 
T. cynrocephalis where 14 mects the lower 
canine, The fossil tooth is wider relative Lo its 
length than are the canines of the dingo (Cunis 
fainiliaris Linnaeus, 1758, var. dingo Blumen- 
bach, 1780). its enamel is smooth rather than 
crenulated .as in phocids, and it is more robust 
than canines of Sarcophilis harrisii (Boitard, 
1841). T have not examined any §. lantarus 
(Owen, 1838). 


S.A.M, P161206 is &n incisor, probably a 
tight 14, 3.9 mm wide, 4.3 mm Jong and with 
a crown height of 4.6 mm. The root is robust, 
curved arid entered the premaxilla to a depth of 
12.9 mm. The occlusal surface is in two planes. 
the larger surface being the plane of wear 
against 1%, the smaller heing the contact sur- 
face with the lower canine. The fossil tooth 
lacks the Jateral cuspules found in incisors af 
c.f, dingo, lacks the transverse groove of pho- 
cids and is more cuboid than the incisors of 8. 
frarnisil, where the incisors are crowded and 
compressed laterally. 


A second incisor tooth, §.A.M. PL6120j, is 
probably # right Ij. It is 4.2 mm wide, 5.1 
mm long and the crown is 3,9 mm higi, The 
root is deep and straight; the tip has been lust. 
The wear surface is fnceted in Wyo planes, the 
Jarger being the wear surface against [4, the 
smaller that against T%. In neither C, f. dingo 
nor phocids are the lower incisors faceted, and 
in §. harrisil the lower incisors are compressed 
like the uppers, 


Additional teeth of T. cyrocéphalus were 
found associated with remains of a Protemne- 
don (c.f, P, brehus) in a rock pile a short 
distance {rom the silt deposit. These teeth were 
almost certainly derived from one indtyidual, 
aod comprise two upper canines, a left mandi- 
bular canine, six of the eight upper incisors and 
all six lower incisors All are comparable in 
size to those of modern adult male T, cyro- 
eeplelas in the South Australian Museum but 
are larger than those of modem adult femates, 


130 


(1. cynocephalus is strongly sexually dimorphic 
(Ride 1964).) 

T. cynocephalus has heen extinct on main- 
land Australia since before European scttle- 
ment, although it existed then in Tasmania. 
During the Pleistocene, however, it was wide- 
spread on the mainland. as shown by its re- 
mains jn cave deposits in Victoria, New South 
Wales, South Australia and south western Aus- 
tralia (Ride 1964). 


Family DASYURIDAE 
Dasyurus maculatus (Kerr, 1792) 

The only identified fragment of D. maculatus 
is a broken Icft maxilla (S.:A.M. P161151) 
containing the canine ulveolus and the six cheek 
teeth, These do not differ in size or morphology 
from those of modern specimens, Some tooth 
dimensions of the fossil are: Pl, length 3.3 
mm, width 1.8 mm; P?#, 14.4 mm, w 2.5 mm; 
M1, buccal length 6,1 mm, w 4.3 mm; M2, 
bi 6.6 mm, w 5.3 mm; M3, bl 6.6 mm, w 
6.6 mm; M4, bl 1.6 mm, w 7.2 mm; M1-3, 
1 (9.5 mm. 

D. maculatus was not rare in the south-east 
of South Australia early in this century (Jones 
1923, p. 88) but is now extinct in this slate 
(Aitken 1970). 


Dasyurus viverrinus (Shaw, 1800) 

The skull of D. viverrinus can be distin- 
guished from tbat of the similar-sized D. geof- 
froii Gould, 1841, by the posterior palatal 
vacuities which ure small in viverrinus but large 


MEREDITH J. SMITH 


in geoffroii (Thomas 1888). The posterior 
palate is not preserved in any Victoria Cave 
specimen. 


The teeth of specimens of the two species 
in the South Australian Museum, and of Vic- 
toria Cave specimens, are similar in size and 
morphology (Fig. 4), and there is overlap in 
all linear dimensions of individual teeth and of 
toothrows. The ratio of the distance from pro- 
tocone to unterior stylar cusp, to the distance 
from protocone to posterior stylar cusp, is 
significantly greater in Ml and M% of D. 
seoffroii Uban in D, viverrinas. (M1! D. viver- 
rinus, Tatio — 0.525, D. geoffreii, 0.570, P<:05 
(t test)! M3: D, viverrinus, 0,659, D. geoffroii, 
0.634, not significant; M%: D,  viverrinus, 
0.664, D. geoffroii, 0.733, P<.05 (t test) ). The 
ratios in the Victoria Cave specimens arc closer 
to those of D. viverrinns (Victoria Cave MV 
ratio, 0.507, M2, 0.623, M3, 0,486) and be- 
cause of this similarity, the Victorian Cave 
specimens are referred to D_ viverrinuy. 
Examination of more complete material could 
possibly alter this. decision. 


Twenty-one maxillary fragments and 34 
mandibular fragments were recovered. Most of 
the latter lacked teeth and although many 
isolated tecth were found, none could be litted 
fo any particular jaw with certainty, Twenty- 
two of the fragments were from adults, 33 
from juveniles, A minimum of seyen adults and 
12 juveniles are represented. Some dimensions 
of the teeth are given in Table 6. 


TABLE 6 


Same dimensions of teeth and alveoli of Dasyurus viverrinus frant Viewria Cave. 


Dimension n Range (mm) 

Mi length 3 5.0—5.7 
M2? length 6 5.0—5.5 
M4 length 2 §$:3—5,4 
M34 length 1 1.2 
M1 alveolar length 4 14.5—45.1 
P4. alveolar length 4 3,1—4.0 
Mj alveolar Jength 10 3.94.9 
My; alveolar Jength 10 4.2—5.0 
M¢ alveolar fength 14 4,0—5,1 
Mj alvealar length 5 4.3—5,3 
M44 alveolar length 4 18.0—20.1 


Standard Coefficient of 

Menon error Variation 
§.30 208 68 
SAS 76 Mh 
5.33 50 13 
14.68 152 2.1 

3.4 196 Ws 

4.25 ALI 6.2 
453 L074 5.2 
4Ad O71 6.0 
4.72, 166 738 
19.10 528 5.5 


SMALL FOSSIL VERTERRATFES FROM NARACOORTE IL 


D. viverrinuy is found in mary cave deposits 
in Western Victoria. (Wakefield 1964) and has 
heen found in the Bat Cave, Naracoorte (Tide- 
mann 1967). The species was formerly com- 
mon in South Australia (Jones 1923, p. 91) 
but it is now extinct in this state (Aitken 1970). 


Genus ANTECHINUS 


Specimens of Antechinus were distinguished 
bv the following criteria. 

(1) The maxillary molars are more robust, 
and less. compressed antero-posteriorly 
than in Sminthopsts. 

The mandibular fourth prémolar is ve- 
duced and is always smaller than P&, 
in Syrunthopsis and <Antechi- 


Gi) 


whereas. in 

nomys P4 is larger than P4, 

Generally the mandible is more robust 

than im Srinthepsis and the masseteric 

fossa wider. However, some small man- 
dibles of A. stuartii are similar in size 
to those of large S, murina, 

Q¥) The entoconid is always well-developed 
as it is in Sminthopsis eraysieaudata. Yt 
is much reduced or absent in other 
species of Sminthapsiy and m Antechi- 
nomys (Bensley 1903). 


rit 


Antechinus flavipes (Waterhouse, 1838) 


The mandibular molars of modern speci- 
mens of A, flavipes in the South Australian 
Museum are robust, the average width of Mi 
being 1.34 nim and that of Mg, 1.42 mm 
(Table 7), The length of M4—4 is equal to or 
greater than 7,2 mm and the length of M1—* 
equal to or greater than 5.5 mm. The premolar 
teeth are broad and crowded, leaving no spaces: 
between adjagent teeth (Fig, 5). 


34 


From Victoria Cave, 10 maxillary and 40 
mandibular fragments from a minimum of 23 
individuals were indistinguishable in mor- 
phology and size trom those of the modern 
specimens of A. flavipes (Table 7). All were 
adults, I have nor examined skulls ef Phasco- 
pole calura Gould, 1844 and from published 
descriptions I cannot exclude the possibility 
that some of the Victoria Cave mandibles are 
of that species, 

Antechinus flavipes inhabits rainforest. dry 
sclcrophyll forest and woodland, where the 
animals obtain their insect food from the tree- 
trunks and large limbs, and from logs. Isolated 
populations are found in north-eastern Queens- 
Jand and in south-western Western Australia, 
while the main population ranges from south- 
eastern Queensland through eastern New South 
Wales to Victoria and south-eastern Australta, 
its distribution being mainly on the inland side 
of the Great Dividing Range, but extending to 
the coast at both the northern and souwth- 
western extremities (Wakefield & Warneke 
1967). Naracoorte is within this range, Re- 
mains Of A. flavipes have been found in the 
Wombeyan Caves, New South Wales, in uv 
deposit that. is probably Upper Pleistocene in 
age (Ride 1960), but have not been found 
in Pleistocene (nor Recent) layers of 
McEachern’s Cave, in the extreme south-west 
of Victoria (Wakefield 1967). 


Antechinus stuartii Macleay, 1841 

The dentition of A. shenrff is identical mor- 
phologically with that of A. flevines and, 
although the former species is on the average 
much smaller, there is overlap in all dimensions 
of skull and teeth (Wakefield & Warneke 


TABLE 7 


Camparlyonys of some dimensions of teeth and mandibles of Antechinus flavipes frem Victoria Cave with 
thase af a madern sample from southern South Aastralia and south-western Victoria. 


A. flavipes from Victoria Cave 


Moder A. flavipes (mn — LS] 


Dimension Standard Standard 
a Range (mm) Meun error CL, Range (mm) Mean error CV- 

M12 alveolar 

length 5.9—6. i 3.98 O31 13 43.3—6.9 f,12 O87 6.0 
M's width 5 2.124 2.28 058 5.7 2,1—2.6 2.36 30 55 
Length of ascend- 

ing ramus 4,5—5,9 $5.1] 96 $.2 45—S5.7 5.19 O77 63 
Ma width 16 12—1.4 1.342 NG 0 1.2—1A 1.34 O17 $2 
Ms width 23 1.3—Ls 1.37 O12 44 1,3—1.5 1,42 19 55 
Mj alveolar 

Jength 23 1.5—1.9 1.75 .022 543 1,8—2.4) 1.88 As 4,3 
M4—4 alveolar 

length 2] 7.28.1 7.354 49 3.0 7.2—8:1 7.59 2065 3.7 


132 


1967). After mensuring modern specimens of 
both species I have arbitrarily chosen to dis- 
tinguish as stuartii all specinrens in which the 
alveolar Jength of M3—% is equal 10 of less Lhun 
5.7 mm and that of M{—, is equal to or less 
thun 7.1 mm, In both modern and fossil man- 
dibles the premolurs wre markedly crowded 
(Fig. 6), with P4 often being sct obliquely to 
the line of the jaw. 

Six maxillary and 16 mandibular fragments, 
from a minimum of IL animals, have been 
found in Victoria Cave. All but one are adults. 
They are similac in morphology and size ta 
modern specimens (‘Table 8). 

A. ytnartit has not heen recorded previously 
from South Australia, although its present 
range extends us far west as Portland in Vic- 
toria, only 70 km east of the South Australian 
border (Wakefield & Wurneke 1967). Its re- 
mains ave common in cave deposits in western 
Victoria where in MecEachern’s Cave in the 
extieme south-west, it is found in both Pleisto- 
cene (15,200 -© 320 years BP) and Recent 
layers (Wakefield 1964, 1967). 

The present ranges of A. stuartii and A. 
flavipey are complementary, the distribution of 
stuarué being coastal to that of flavipes, but 
overlap docs occur, e.g. in western Victorin 
(Wakefield & Warneke 1967). 


Antechinus swainsonii (Waterhouse, 1840) 

A, swainsenii and A. minimus (Geoffroy, 
1803) ale characterized by their long claws 
and jong snouts, The molar teeth are as. long 
as, or nearly as long as, those of 4, fluvipes 
e.g. 1 3 specimens of A. sweinvenii (S.A.M, 
M2421, M7047 and M7496), M1-3 — 5.5- 


MEREDITH J. SMITII 


6.0 mm (mean 5.73), M4-4 = 7,527.8 mm 
(7.6)], but ate much narrower [M8 = 1,9- 
2.0 mm (1,97), Mg = 1.1 mm (in all 3 
specimens), M4 — 1,1-1.2 mm (1.17)], The 
premolars too are much narrower than in A. 
flavipes and are: not crowded, «adjacent teeth 
often being separated by a space. In addition, 
the mandibular premolars have long talonids 
with sharp posterior cusprles, whereas the 
taloitids of A. flavipes premolars are short with 
blunt cuspules. The mandibles of 4. swainsenti 
and 4. minimus are more slender than those 
of A. flavipes, and longer than those of Syin- 
thepils (Fig. 7). 

Nane mandibles, from a minimum of five 
animuls, conformed with the swaisenit-ninl- 
mas characteristics, and I have tentatively 
classified them as A. swainvonii because all 
have uw long mandibular symphysis, extending 
posterior to the front to PY. “lhe symphysis 
In A, snininuix is shorter (Tate 1947). Tr 
addition, the preatest breadth of the masseteric 
fossa in four Victoria. Cave specimens ranges 
from 4.2 to 5.0 mm, whereas in A. meinirnus 
its greatest breadth does not. exceed 4.2 mm 
(Thomas L888). In Victoria Cave specimens, 
the mean width of Mi is 1.13 mm (3 spee,), 
mean width M4 is 1.22 mm (4 spec.) and 
length Mq— ranges from 6.7 to 7-2 mm (meyn 
7,0) in four specimens. 


A. swainseni? has not been recorded ulive 
in South Australia (Aitken 1970), but its re- 
mains were found in a late Recent deposit in 
the Bat Cave, Naracoorte (Tidemann 1967), A 
single, incomplete, toothless mandible from 
level | of the Frommy’s Landing archaeological 
excavation on the River Murray was assigned 


TABLE 8 


Cempurisons of some dimensiony of teeth and mandibles of Antechinus stuartii fram Vietoria Cave with 
ioxe of a modern sample from Bondo, NSW, 


4, strartii from Victoria Cave 


Modern A. stuartil (on — 10) 


Dimension Standard Standard 
nm Rangetnim) Mean erro CV. Range({mm) Mean error CY, 

Mi—% alveolar 

Jengih a §.1—5.7 5.50 141 5A 5335.9 5.57 N68 3.9 
M3 width a 2.0—2.:2 2.43 AGT 5.4 1.9—2.1 1.96 .022 3.6 
Leweth of ascend- 5 42—5,4 4,40 207 1.1 3.9—4.4 419 083 3.Z 

ing Tums n 7 
Ma width S 1-13 1,22 .037 69 1i—.3 120 OLS 

h Width 7 126-13 |.24 a0 43 1,213 1,27 O15 a8 
M4 alveular ‘ . 

length 1d 1.4—1.3 1,72 025 46 15—1.8 1,69 038 71 
M4-4 alveolar 

lenzth T S3—7.1 f.AT 180 72 6,5—7,0 4.85 052 2.4 


SMALI. FOSSIL VERTEBRATES FROM NARACOORTE II 


to this species (Wakefield #2 Mulvaney ef al, 
1954), 4, 8weinsenii is commonly found in 
cave deposits 19 Western Victoria (¢,g- Wake- 
ficld 1964, 1967), On the Australian mainland, 
A. ominimay bas a limited range around the 
South Australian-Victorian border near the 
coast (Wakefield & Warneke 1963). 


Genus SMINTHOPSIS 


Fragments of Siminthopsis were identified by 
the relatively Jarge P4 and by the greater antero- 
posterior compression of the maxillary molars 
than in Antechinus. The mandible is generally 
more slender than m <Anlechinus and the 
ascending ramus shorter antero-posteriorly, but 
there i¢ overlap between S. murina and A. 
siucré in mandible size, 


Sminthopsis murina (Waterhouse, 1838) 


Mandibles of S. murine may be distinguished 
from those af S_ crussicaudata (Gould. 1844) 
hy differences in the morphology of the 
talonrds, The entoconids are reduced or wbsent 
in S. marina but well-developed in 3. cressi- 
caudata (see Bensley 1903}. In the maxillae, 
interdental fenestrae are smaller and less 
numerous in §. purina than in S_ crassicaudata 
(pers. comm. Michael Archer, Western Aus- 
tralian Museum), 


In Antechinomys luniger (Gould, 1856), 
which also lacks the entoconid, the postero- 
external shelf of the lower molars is much 
broader than in S. murina, The dentition of 
S. leucopus is said to be distinguishable from 
S. murina by the presence of spaces between 
adjacent premolar Ieeth (Thomas 1888), but 
this character is yariable in the specimens of 
S. mvrina in the South Australian Museum. 


1s 


io 


Sixteen maxillary and 57 tooth-bearing man- 
dibular fragmenis from a minimum of 31 
animals were found; Only two were juveniles, 
A further 61 toothless mandibles. 32 right and 
29 left, are probably referable to this species. 
Some dimensions of the adult specimens and 
of a modern sumple are given in Table 9. The 
maxillary interdental fenestrae occupied a 
larger proportion of che interdental space than 
In many modern specimens and the entoconid 
Was not present i anv Victor Cave mandible. 
Adjacent premolars, both maxillary and man- 
dibular, usually touched; there was never a 
conspicuous gap between the premolars (Fig- 
R). 

Tn addition to the specimens. listed above, 
one small adult mandible (S.A.M, P16118z) 
was found that is morphologically identical 
with S. murina but is much smaller, the length 
of M4—{ being only 4.8 mm, 

5. murina is widespread in South Austratia 
but ig nowhere common (Jones 1925, p. 118, 
Aitken 1970). There is a specimen in the 
South Australian Museum from Bordertown 
but Tidemann (1967) did not find this species 
in alate Recent deposit in the Bat Cave, Nara- 
coorte. The morphologically-similar species, 8. 
leucopus (Gray, 1842), is found in Recent, 
but not Pleistocene layers in McEachern's 
Cave, extreme south-western Victoria and in 
other cave deposits in south-western Victoria 
{Wakefield 1964, 1967), 


Sminthopsis crassicandata (Gould, 1844) 

S. crassicaudeta was tepresented by only 
three mandibles and one maxilla from a mini 
mum of three individuals, all adult, Some 
dimensions of these are; length of M1-—4, 4.8 


TABLE 9 


Comparisons of some dimensions of teeth and mandibles of Sminthopsis murina from Victoria Cave 
with those of a modern sample from South Anstralia. 


_S. muritia from Victoria Cave 


Modern §. murine (n— 10) 


Dimension Standard Standard 
n Ranzgetmm) Mean ermr = C.Y. Range (mm) Mean error CY, 

Mi—} alveolar : 

length 46—5,0 AT4 050 3.2 4.5—5,2 4.76 D6z 4 
M3 width 9 19—2) 1.98 (722 3.4 1.7—2.0 1.85 27 4.6 
Length of ascend- 

ing ramus 5 3.5—4.A 4.06 $77 7.3 3.44.6 4.05 137) (107 
M4 width 7 1.0—1.1 1.os O42 49 o.9—1,1 1,00 ALS 4.7 
M4 width 23 iI—12 L.14 O10 4.4 io 2.2 1.09 £028 8.0 
M4 alvensar 

length 3a) 1.41.7 1.48 014 5.2 1414 1.56 037 TA 
M4, alveolar 

Iength 26 5.4—.3 5.598 034 3.6 § 54.2 5.88 AT 4,2 


a 
& 


mm; length of ascending ramus (3 specimens). 
3,.6-3,9 mm (mean 3.73); M4 width (3), 1.0 
min, My alveolar length (3), 1.4015 mm 
(1.43); alveolar length M4—, (3), 5.5-5.7 mm 
(5.67). 


This species has w Wide range in southern 
Western Australia, South Australis, Victoria, 
Western New South Wales and south-western 
Queensiand (Ride 1970). and in the South 
Australian Museum there are many specimens 
from the sourf-east of Sopth Australia, Tt was 
found in the Bat Cave deposit, Naracoorte 
(Tidemann 1967) and in Recent cave deposits 
ity sQuth-weslern Victoria (Wakeficla 1464, 
T2847), 


Discussion 
Method vf accumulation of the small vertebrate 
remains, 

Analysis of the remains of cach species, intu 
udults and juveniles, shows that there is con- 
siderable vatialion between species Ih the pro- 
portion of adults. The larger species, Betforgia 
penicillata and Petorous apicalis, are represen- 
ted almost entirely by juveniles (Smith 1971), 
as is the large bundicoot, Perqmeles eutictit. 
Adults and juveniles of smaller species, such as 
Isoodon ebesulus and Potorous platyops, were 
found im about equal numbers. while the small 
dasyurids Canrechinus and Saaethopsts) ind 
the petaurids and burramyids were nearly all 
adults. Slower eruption of the teeth in the 
larger species may account for some of this 
variation, but it dogs not account for the wide 
vatiations scen between two species of the one 
fenus. For cxample, within Pevoneles no more 
than one qiarter of the larger P. gunaiz were 
adults. but more than half of the smaller P, 
fouguinville were adults: 


The |iased age structure suggesls chat the 
cave unl mut act as a simple pitfall trap. but 
thal the bones were brought in by precuturs 
able to capture animals as large us an aduit 
P. bougainville or FPetorous platyops, or a 
juvenile P, yunnii or bettony. Mammal predu- 
tofs thik inhabil dens usually dic within them 


MPREDELH J. SMITH 


occasionally. Thy facies cynecephalus was able 
lo take larger prey (Ride 1970) and so the 
predators could have been dasyurids, Datyuris 
ynucwlatus or D. viverrinus, or owls, The very 
low incidence of D. maculatus sugeests that 
this species was nut the predator. D. viverrinus 
is hetter represented but the high proportion of 
juveniles indicates Jhat it was a prey species 
father than a predator. The small manuals 
therefore probably accumulated from owl pel- 
lets. The method of accumulation of the Jarec 
herbivores in the deposit is mnt yet known, 


Climatic interpretapions 


Modern populations of the scansorial species 
Antechinus flavipes and A. stuartii are sympat- 
ne in areas of dry sclerophyl Forest such us 
ut Glenlofty, western Victoria, where strinay- 
bark (Bucalypries inecroretiyachiy «and box te. 
melfiodara and &. goniocatyx) are usyociuted 
with a sparse ground cover of sawsedge 
(Guhnia vadula) and tussock grass 4 Porc 
(Wakefield & Warneke 1967). Populitions of 
the ground-dwelling A. swainsenii are densest 
in wet selerophyll forest, bur they also occur in 
other hahitais such as open woodland and 
stunted coastal euealypt scrub with tussock 
grass (Wakefield & Warneke 1963), 

Modern Smiinthepyiy murina and §. craxsi- 
caidats inhabit hath wet and arid areus and 
little is known of their habitat requirements, 
Peramoles bovguinville was similarly wie 
ranging al the heyinning of European explora- 
tion, and its habitat requirements are also un- 
known, 


P. gman? and 2. odexuluy oecur sympatrically 
in Tasmania, Both species require s¢rub tor 
nesting and the food of both consists mainly of 
eathworms and insect durvae, However. 
Iseodon remains within the scrub to feed, 
Whereas Permmeles forages Car out into open 
areas (Heinsohn 1966). If the Pleistocerie 
Peranteles and fsoadon had similar ecological 
requirements to their modem descendants, one 
might infer whet the apparent scarcity of 
fyooden indicated that the vegetation of the 
atea was an open woodland, with littl dense 
scrub. On the other hand, the greater abun- 


Lett maxilla of Dasyurus viverrinus, (S.A,M, P16L15a) from Victoria Cave. occlusal view of 


Fig, |. Left mandible of fsoaden wbesulus (S.:A.M. P16112%) from Victoria Cave. 
Fig. 2. Right mundihle of Perameles gunnii (S,A0M. P1610dv) from Victoria Cave 
Vig. 3, Teft mandible of Perameles houvgainville (S.A.M, P16103f) front Victoria Cave. 
Fig. + 

M!, lo Mt. 
Fig, 3, Right mandible of Antechinus flavipes (S.A.M. P16001)) from Vicloria Cave. 
Fig. f. Right mandible of Antechiaus stuarti (S:A.M, P16119) from Victoria Cave. 
Fig- 7. Right mandible of datechinus swainsenii (S.A.M. PI60D9a) fran Victoria Cave. 
Fig, 8 


Right mandible of Sminthopsis murina (S.A.M. P16021a) from Victoria Cave, 


SMALL FOSSIL VERTEBRATES FROM NARACOORTE II 


we 
wn 


136 


danee of Peraneles might be wn artefact of 
selective predation, for unimals foraging in 
scruh would be less susceptible to owl attack 
than animals foraging in the open, However 
in the Mammoth Cave deposit, P, bougalniville 
js about twice as abundant as 7. obesulus. This 
in believed to be a true reflection of a larger 
population of Perameles in the Pleistocene, for 
the deposit does nol scent to have originated 
from owl pellets, but appears to have been a 
talus deposit, uccuomilating as animals tell 
through holes in (he roof (Merrilees 1965), 


The combined evidence of the represented 
species of potoroines, petaurids and burramyids 
(Smith 1971) and of the perumelids and 
dasyurids indicates that at che time of accumu- 
lation of the deposit. Victoria Cave was sure 
rounded by dry scleraphyll forest 


MEREDITH J, SMETH 


Acknowledgements 

Excavation of the deposit would nat have 
proceeded without the enthusiastic help of 
CEGSA members in divging and sieving 
Transport costs for these helpers were delrayed 
by a grant from the South Australian Goyern- 
ment Tourist Bureau. The late Mr. B. Maddock 
ubly mediated between CEGSA and the Tourist 
Bureau and actively assisted in the working ol 
the deposit. 

Tam grateful to Mr. P. F. Aitken, Mr. M, 
Archer and Mr, R. TF. Wells for many helptul 
discussions and for their eviticism of the manu- 
script. To Mr, Archer fF am especially grateful 
for his dvice on the separation of species of 
Antechinus and Sminthopyixs, The photographs 
were prepared by the Photography Department. 
University of New Rngland (Figs, l3, 7) and 
Mr. E. Sangster (Figs 6 and 4). 


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371-391. 


137 


WAKEFIELD, N. A. (1966b).—Mammals recorded 
for the mallee, Victoria. Proc. R. Soc. Vict. 
79 (2), 627-636. 


WAKEFIELD, N. A. (1967).—Preliminary report on 
McEachern’s Cave, 8,W. Victoria. Vict. Nat. 


84, 363-383. 


WAKEFIELD, N. A., & WARNEKE, R. M. (1963).— 
Some revision in Antechinus (Marsupialia) I. 
Vict. Nat. 80 (7), 194-219. 


WAKEFIELD, N, A., & WaARNEKE, R. M. (1967).— 
Some revision in Anfechinus (Marsupialia) 2. 
Vic. Nat. 84 (3), 69-99. 


THE GENUS ACUARIA BREMSER (NEMATODA: SPIRURIDA) IN 
AUSTRALIA 


BY PATRICIA M. MAWSON 


Summary 


This paper lists all the known Australian species of the genus Acuaria (sens. str.). The degree of 
infestation in families of passerine birds is indicated in a table. New species described include A. 
petterae of which males, with or without females, are recorded from Lalage leucomela (type host), 
Meliphaga virescens, M. plumula, Cracticus nigrogularis, Artamus melanops, Cinclosoma 
cinnamomeum, Myiagra inquieta, and Drymodes brunneopygia, and females, probably of this 
species, from Acanthogenys rufogularis, Anthochaera carunculata and Oreoica gutturalis. Other 
new species are Acuaria colluricinclae from Colluricincla rufiventris; A. microecae from Microeca 
leucophaea; and A. mirafrae from Mirafra javanica. Measurements and some redescription are 
given of A. anthuris from Corvus melanops, C. coronoides, C. bennetti and C. orru; A. streperina 
from Strepera versicolor; and A. skrjabini from introduced aviary finches, Tiaris canora, Lonclzura 
malacca and Estrilda melpoda. 

Characters considered useful in distinguishing species of this genus are cordon length and pattern, 
the shape and ratio of the lengths of the spicules, and the number and arrangement of the caudal 
papillae of the male. A key to most of the known species, based on male characters, is also given. 


THE GENUS ACUARIA BREMSER (NEMATODA;: SPIRURIDA) IN AUSTRALIA 
by Patricta M. Mawson* 


Summary 


Vhis paper lists all the known Australian species. of the genus devaria (sens. str.). The degree of 
infestation in families of passerine birds is indicated in a table. New species described include A.. 
penerué of which males, with or without females, are recorded from Lalage leucermela (type hest), 
Meliphaga virescens, M, pliemula, Cracticus nigrogularis, Artamus melanops, Cinclosama cinramenmeum, 
Myingra ingaiete, and Drymodes briummeapygia, and females, probably of this species, from Acantho- 
genys rufogularis, Anthochaera carnnenlata and Orevica putturalis. Other new species are Acvaria 
omMlluricinclae from Colluricincla rufiveniris; A. microecag fram Microeca letcophaea; and A, mirafrae 
from Mirafra javanica. Measirements and some redescription are given of A. anthuris from Carvas 
melanops, C. corondidés, C. bennetti and C. orru; A. streperina from Strepera versicolor; and A, skrja- 
bint from introduced aviary finches, Tiaris cunerg, Lonchura malacea and Esirilda melpeda. 

Characters considered useful in distinguishing species of this genus are cordon [cngth and. pat- 
tern, Ue sliape and ratio of the lengths of the spicules. and the number and arrangement of the 
caudal papillae of the male. A key to most of the known species, based on male characters, is also 


given. 


Introduction 
Almost all known species of the genus 
Acuqria Bremser (sens. str.) are from passerine 
birds of the order Oscines; there appears to be 


TABLE 1 
Incidence of Acuaria spy, and af nemaiodes 
generally, ia “land hirds’ dissected, Numbers refer 
10 ypecimens, Not species, 


‘ ? Number With With 
only one exception to this: A. wpupa Rasheed, Bird group dissected = nemittodes Acuaria sp. 
1960, from the coraciiform bird Upupa epops — Passeriformes 958 3a 7 
from India. Acuaria spp. have been recorded plantidae sid Z i 2 
from galliform and gruiform birds, and from fade OR . 4 
cormorants, herons and birds of prey, but ail Monurchidwe 4 7 { 
specics, Of which the mule is described, are Muscicapidae 41 W 1 
found to belong to related acuariid genera. Puchybeyialidpe fs s 2 
Where only the female is described, identifica- ACOneW ese J 
. . : 5 Meliphugidae Ja9 45 4 
tion of the genus is uncertain, but may be Attamidac 10 a) 9 
inferred from the cordon struciure. if this is Cracticidae 100 53 3 
described, Carvidae 77 69 50 

hie tuidittabe apatite Catan. s i" Other families AR? 137 — 

_ The incidence of Acuaria (sens, str. ) Species Cunrimulgiformes 18 Yl _ 
in birds dissected in this department is shown Coraciiformes 28 14 — 
in Table 1. Crows are by far the most com-  Strigeiformes 25 17 -- 
monly infected and are also the most heavily Accipitriformes 61 38 = 
infested birds, perhaps howeyer, only because Caculifocirics <] é = 
TREESTE + Pt P oie ee! - saat Columbiformes 43 1 — 
of the greater size of the gizzard. Of the 21 Psiitaciformes 157 4 = 
smaller passerines listed, belonging to 14 spe- Galliformes 7 va — 
7 a= 


cies, none yielded more than three specimens, 
and eight birds contained only females. Under 
these conditions (and these apparently pertain 
also in other places—-see Chabaud & Petter 
1961), it is almost impossible to be certain of 
the variation within a species. However, in the 
present maternal, two species are present in 


Gruitormes 56 1 


some numbers. 4. aathuris from Corvus spp., 
and A, skrjabini from imported finches (cage- 
birds} among which a heavy infestation 
occurred. Within each of these species there is 
a close agreement in certain characters; the 


* Zoology Department, University of Adelaide, S. Aust 5000. 


Trans. R. Soc. S$. Aust. Vol. 96, Part 3, 31 Angust 1972, 


140 


cordon lengths in male and female (different 
in the two sexes), the shape, size and leneth 
ritio of the (wo spicules, and the number antl 
arrangement of the caudal papillae of the male. 

Speciinens Crom other Australian hosts were 
vrouped together according to these characters; 
m an allempt to compare them with species 
already described, a key to most of the known 
species, based on these characters, was vom- 
piled. ‘Ihis is. given below. 


Examination of the shapes of the cuticular 
bosses in the cordons of the Australian species 
shows that these may be useful in comparing 
them. The detailed structure of the cordons, 
especially us seen in transverse section, has 
becn suggested by Skrjabn et af (1949) as a 
useful genéri¢ character in the Acuariidae, Wil- 
liams (1930) and Rasheed (1960) give figures 
of the surface pattern in some species (though 
those tn the latter publication are too much 
reduced to be of critical value). The patterns 
in each of the Australian species are similar in 
all specimens, of both sexes. 


Acuarta species from Australian birds 


Alaudidac 
MIRAFRA JAVANICA  Horsfielil. 
mircfrae n. Sp. 

Campephagidae 
LALAGE. LEUCOMELA Vig. & Hos, 4, pet- 
terde Ty Sp. 

Turdidae ; 
DRYMODES BRUNNEGPYCLA Gould. A, 
pelferde Nh. Sp, 

Monarchidise: 
MYIAGKA INQUIETA (Latham). A. pedterae 
Th. Sp. 

Falounculidae 
OREOICA GUTTURATIS (Viz. & Hors.), f 
petterae n. 3p. 

Moliphagidae 
MELIPHAGA VIRESCENS Vicillot. 
tart 1. Sp. 
M. PLUMULA Gould, A. petterac n. sp, 
ACANTHAGENYS RUFOGULARIS Gould 
AL petterae 1. Sp. 

Ayiamidae. 
ARTAMUS CINEREUS Vicillot 
fl. sp. 

Pachvcephalidae 
COLLURICINCLA HARMONICA WHITE! 
Mathews. A, calfuricinclie t. sp. 

Cracticidae 
CRACTICUS NIGROGUILARIS (Gauld). 4- 


petterae Tn. s 
(Latham). 4. 


Acuaria 


A. pet 


A. pelierae 


NTREPERA iy VERSICOLOR 
streperina Johnston & Mawson 
Muscicapidae 
MICROECA LEUCOPHARA (Latham). 4. 
microcear N. Sp. 
Corvidoe 
CORVUS CGRONGIDES Vig, & Hors. 4. 
anthnds (Rud) 


PATRICIA M, MAWSON 


C. MELLORI Mathews. A. anthuris (Rud.) 
©. BENNETT? North. 4. anthuriy CRud.> 
C, GRRU Booapurie. A. anthuriv (Rud 


Key for identification of male specimens nf 
Acuaria spp. 

The descriptions of A, gaywnyiy Bisseru and 
A, iwashkini Brhardoya are not available to 
mes u full description has not been seen of 4. 
eremophila Erkulov. A. denuiy Duj. has been 
omitted because the cordon length and the 
number and arrangeroent of the caudal pupillac 
are not known, it falls among species below 
choice 14 in the key, Species from crows, 4 
altenmeta (Rad.j}, Al oernata (Gendreb, A. 
longicaudata Hoeppli & Hsti, and A, seutara 
Maplestone, und synonyms of these, have becn 
assigned to ane group, the “A. anthuriy come 
plex”. Ic is probable that examination of the 
Lypes of all deseribed species attributed to 
Acwearia would show considerable synanymiy, 
unl might also indicate more important differ- 
ences helween some species than ure revealed 
by existing descriptions. 


J. Left spicule longer Iban 190 em. on yD 
1, Teft spicule shotter than 190 em 9 
2. Spicule ratio L.1-14 3 
2. Spicule ratio 1.5 ar more ees 
3, Cordons yery long, extending well past o¢sa- 
phagus “A anthuris Complex" 
3. Paatons very ‘short, not t extending much past 
excrolory pore ov... 4 

4, Body length 4-6 mm, spicule ratip 1.4 
A. muyvori Lem, Freitas & Proenga 
4, Rody Jengih 10-11 mm, spicule ratia 1.1- 
14, _ A, vordata (Mueller) 
S. Teft spiciile Texs. than 230 vim long & 
5S. Lett spicule more than 250 em long 7 
6. Cordons end ubout midlength of muscular 
cesophagis A. subula (Duj,) 

6. Cordons nearly as long as oesophagus 
A. colluricinelae 0, sp. 
7. Lett spioule 262 “im A. turdi (Wang) 
7. Left spicule over 300 am... x 
& First pair of postanal papillae ubutil 2 third 

tail length fronr second pair 

A. cyanavilla (Boyd) 
6. First and second pairs of postanal papillae 
not much separated 4. streperina J..& M 
9. Spicule ratio 1.5 or over 10 
9. Spicule ratio Jess than 1.5... 0... 14 


1). Cordons reach to end of muscular oeso- 


phagus a al, conica Maplestoene 

LU, Cordons very shorl, not much past exerclory 
pore... tl 

11. Six pnirs. of postanal papillae ... 12 
11, Seven pairs of postanal papillae ... .. ... 13 


12 Left spicule 165 wan long ., 

A. sialia Williams 
12. Left spreule 150 “em long 

A. puyittitfera ‘Lins, 


ACUARIA (NEMATODA: SPIRURIDA] IN AUSTRALIA 14} 


13, Left spicule 140 #m long . 
A. paragalliardi Ch, & P, 
13. Left spicule 170 4m Jong . 
A, parorioli Ch & PB. 
14. Cordons more or less to end of glandular 
oesophagus 1s 
14. Gurdon harelly longer than rouycular oes 
phagus _. a 
(§ Four pairs of preanal papillae aes n 
1S. Fewer than four paira of preanal papillae is 
to. Left spicule 129 um long ' 
4. orem) Rasheed 
16, Left spicule longer than 150.am . | 17 
17. Left spicule slightly grooved near Gp 
A. partonl Williams 
17. Left spicule deeply grooved throughout length 
A minor Wiliams 
8. Three pairs of preanal Papillae .. 
A_hrevispicula Maplestone 
18, Two pairs of preanal papillae . . 19 
19, Seven pairs of postanal papillac — - 
A, alii Rasheed 
19. Six pairs of postanal papillae .. 
A. stn Rasheed 


20. Spicule ratio close lo 1.0 . at ap Gs pl 
20, Spicule ratio t4I-14 - - 25 
21. Spicule Jength Jess thaw 130am........, 22 
2), Spicules longer than 130 4m 24 


22. Six pairs of postanal papillve 
A. cremopiila Erkuloy 
22. Seven pairs of preanal papillae in 23 
23, Cordons not much past nerve ring .. 
di. Keng Singh 
23, Cordons reach aboul to end of muscular 
oesophagus _ .,,_ ,_ . A. micraccoe n. sp. 
24, Cordons not past nerve ring ., 
A. martinagliai Le Roux 
24. Cordons nearly to end of muscular deso- 


phacus |, , A upupa Rasheed 
25, Six pairs of postanal papillae... ... 26 
25, Seven pairs. of postanal papillae 31 
26. Postanal papillae in two groups of three 
pairs... _ 27 

26. Postanal papillae not in 1wo distinct 
BrOUPS ae 29 
27. Spicule ratio about |.] A. enter itae mn, sp, 
27, Sptcule ratio |.3-1.4 . 28 


28. Caudal alae widen al. mmidllengti 
A. giisenfe Williams 
28, Caudal alae about same width 
throughout .. ..., A, dolifasi Ch, & Petter 


29, Cordons reach onty to cervical papillae... .., 
A. cettiag Heit 

29. Cordons reach further than cervical 
papillae 0. 30 


30, Right spicule grooved for n most bof its 
Tength A. gracilis (Gendre) 
30. Right spicule ‘simple - A. dierura Rashecd 


31. Three pairs of preanal papillac ... 
A, brumpti Ch, & Petter 
31, Four pairs of preanal papillae 32 


32. Left spicule Jess than 125 4m iaae ‘ 
. A. galliard’ Ch. & Fetter 
32, Left spicule more than 135 #¢m long .,,. 33 


33. End of right spicule enlarged . 
A. skriablad Ozerska 
33, Tip of night spicule without prominent 
enlargement 34 


34. Cordons reach past excretory pore, and 
more than half distance between bead and 
posterior end of muscular oesophagus - 

A. butinerae Ch. & Petter 

34. Cordons shorter, less a half Wis 

Mistames ay _ 35 


35. Parasitic in African oriate ‘ 
A. orioll Ch. & Petter 
35. Parasitic in Australian passerines .... ..., 

A. petterae n. sp. 


Descriptions of Species 

The general morphology of Acuaria spp. is 
so similar that only the special features of each 
species Will be described, Measurements are 
given in Table 2; those of paris of the ocso- 
phagus are taken from the anterior end of the 
body to the end of the organ in question; the 
spicules are meagured in lateral view (often 
very different from those taken in ventral 
view), 


Acuaria anthuris (Rudolphi, 1819} 
FIGS, 1-3 

Hosts and Jocalitics: Corvus coronaides fram 
Adelaide and Pt, Augusta, S, Aust; ©, 
mellort from Balgowan, S. Aust. and Laun- 
ceston, Tas,; C. bennetti from Lock, 8. Aust 
and Erldunda, NT; C. ofre feom Plenty 
River, N.T.; C- sp. from Pearson 1, S. Aust, 


Acuaria cnthuris has been recorded many 
times from different parts of the world; refer- 
ence lists and discussion of its synoymy may be 
found in Skrjabin et al (1965) and Chabaud & 
Petter (1961). The present study deals only 
with the variations observed in the Australian 
specimens. The species is quite common in 
Australian crows and ravens. Measurements 
are given in Table 2. The general appearance, 
except where noted below, agrees with descrip- 
tions given by Singh (1948), Rasheed (1960) 
and Chabaud & Petter (1961). 


The cordons extend well past the oesophagus 
in both sexes, reaching a little under a third 
of the bady length in the male and a little more 
than this in the female, but never quite reach- 
ing to the vulva, The cordon structure (Fig. 1) 
is different from that figured hy Rasheed, 


The papillae on the male tail are usually 
more or less symmetrical, comprising four 
pairs and one median preanal paplilae, and six 
paits of postanal papillac, arranged as three 
pairs on the anterior half of the tail and three 
pairs of rather smaller papillae on the last 


PATRICIA M. MAWSON 


142 


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ACUARIA (NEMATODA: SPIRURIDA) IN AUSTRALIA 143 


quarter of the tail, as well as a pair of yery 
small phasmids almost terminaliy. The mem- 
hers of a postanal pair are not always strictly 
opposite to one another. Individual variations 
from this occur, some specimens having one 
or two papillae missing from one side or the 
other. Of 85 male worms examined, 18 showed 
some abnormality in the caudal papillae. Most 
of these were one papillae more or less on one 
side or the other; in a few there was one papilla 
more or less in the terminal group of postanal 


pupilluc. In three specimens there were six 
pairs in the preanal group, the most posterior 
of these lying just posterior to the anus, so that 
they could be regarded as an extra postanal 
pair except that they continued as a closely 
spaced line of small preanal papillae on each 
side and were quite separated from the larger 
papillae of the anterior group of postanal 
papillae which were further apart. Except for 
these three specimens, all had six pairs of post- 
anal papillae, of which the anteriormest lay 


4 toe 
lo 
=————— 
—— 
3 
\\ 1 nm 
Segk= 
=06c 
| 3o0F 
2 Soo = 
f eq 
7 =3eE8a— 


Figs. 1-3. 

female, 
Figs, 455. 
Figs, 6-9. 


_ 
= 
le 
1 


Soge 
mS == 
——T > 
S03 = 
SHe= 13 
6 1S pint 
_ os a 
5S (c= 
oe 
set 
2 a 
JS( tC 
S34 E= 


Acuaria anthuris. Fig, 1,—Part of a cordon, Fiz, 2.— Right spicule. Fig, 3—Tail of 


A, strepering, Fig. 4.—Part of a cardon. Fig, 5—Posterior end of male. 
A. skrjabimt. Fig. 6.—Part of a cordon. Fig. 7,—Anterior end of male. Fig. §—Pos- 


terior end of mule, Fig. 9—Tail of female, 


Figs. 10-13. 


A petrerge. Fig, 10—Anterior end of male. Fig, 11.—Parl. of a cordon. Fig. 12.— 


: Posterior end of male. Fig. 13.—Tail of femalc. 
Figs. 2, 5, 7, and 13 to scale beside 2; figs, 8, 10, 12, and 13 to scale beside 12. 


144 PATRICTA M. MAWSON 


some distance hebind the anus. ‘The spicules 
are grooved (as described by Singh 1948) and 
alate (Fig. 2), The Jeft spicule is larger thai 
the right except al the lip und the expanded 
pirts of the alac are wider. 

Chabauu & Potter (1961, p. 210) report A. 
anthuris of two types; the first (from Gurritlus 
glandarius and Piew pica), rather smaller, with 
six pairs of postanal papillae in the male; the 
second (from Corvus corone). larger and with 
seven pairs of postanal papillae, in addition te 
the phasmids, The only measurement given is 
(hat the males of the smaller specimens are less 
than 12 mm long. The smaller specimens agree 
with Rudolphi’s specimens selected from 
material (apparently containing more than one 
species), by Schneider (1866) as the type for 
A, anthuriy, Vhe Australian material, though 
perhaps a little longer, agrees wiih these types 


Acuarla streperina Johnston & Mawson, 1941: 
154, 
FIGS. +5 
Host und locality: Sirepera versicolor melan- 
optera from Waikerie, S. Aust, 

The type specimens, of A streperina have 
been re-exuimined and the onginal description 
must now be amended; they are old specimens, 
poorly fixed and much contracted. The length 
given for the oesophagus, 700 ym in the mate 
and 800 ym in the female, is that of the mus- 
cular part of the organ; the end uf the vlandu- 
lar part is 2.1 mm from the head in the female, 
which is strongly contracted, and 1.4 nim in 
the male, which is less so. The cardons reach 
nearly to the end of the oesophagus in the 
Female, andl to the end of the muscular oeso- 
phagus in the male, There are six (not five) 
puirs of posteloacal papillac in the male, 
atringec! with three well spaced pairs on the 
proximal two-thirds of the tail and three pairs. 
closer together, on the distal third. The papillae 
of the fatter yroup are much smaller and 
harder to find. The spicules each have an 
enlarged proximal end, which is less heavily 
ghitinised and was apparently not included in 
the original messurements. The spicules ure 
IO am and 120 um dong, with a ratio of 
1:1.7, The largest eggs are 45 x 28 ym, 

A single female worm from the type host 
species is referred ti A. streperina. [It was col- 
lected and fixed after death and so is in a 
relaxed condition. fis measurements are cilfer- 
ent in those of the type [cmale largely because 
of this, Eges in this specimen are mot em- 
bryonued and are thin-shelled, Measurements 
are given in Table 2.. 


The species is very close to A. cranecitia 
(Boyd, 1950) bat is distinguished hy the 
avangement of the pastanal papillae in the 
male 


Acuaria skrjabini Ozersha, 1926; 103-1117 vide 
Skrjabin v7 ai. 1965: 114. 


FIGS. f--9 


Hasts and locality; Exotic aviary finches from 
New South Wales: Tiuriv canora, Lonchura 
maleces and Kstrilda melpada, 


These specimens oceutfed in large numbers 
in many specimens of the finches and ware con- 
sidered by the owner of the aviary to be the 
cuuse of the death of the birds. They degree 
generally with the figures and description of A. 
skrjabiné by Ozerska and also by Siagh (194874, 
the principal differences being [hat there are 
7 pairs of postanal papillae in the male. as 
described by Singh. mot six ay shown by 
Ocersky; the spicule ratio is nearer that in 
Ozerska’s specimens than those of Singh. Where 
isa distinct enlargement al. the distal end of the 
Tight spicule, 

The comons in the male reach to, and 
usually beyond, the excretory pore, and those 
of the female are longer. reaching to about half 
the distance from the head to the end of the 
muscular oesophagus. 


The caudal alae of the male are distinctly 
wider anteriorly, There is only a slight distine- 
tion in spacing between the first four pust- 
cloacal papillae and the Jast three. In some 
specimens the postcloacul pairs are not 
arranged symmetrically and ina few one mem- 
ber of a pair is absent, Both spicules are in- 
dented at the tips and this is clearer in the right 
spicule as i ends more broadly, 

The cag size is 40-43 by 23-24 ym: this is 
rather shorter than Oxerski’'s measurements, 
and clistinetly larger than thase of Singb. 


Acuaria petterae 1, sp. 
FIGS. 10-13 


Hosts and loculitiess Lalage lencomela Srom 
Katherine Gorge, N T., type host; Meliphuge 
virescens, M, pluntinia and Cractieus nigro- 
eularis from the Pelermann Ranges, N.T.; 
Arianius melanops from Alice Springs, N.T.; 
Cinclesoma cinnummameum trom Tobermory 
Sta., N-Tuy Myiaera inquieta and Drymades 
hrunneopysie from Blanchetown, S, Aust. 

Probable hosts and localities (only females 
present): Acanthogenys rufogularis trom 
Blanchetown, 8. Aust; Anéhochuera carun- 


ACUARIA (NEMATODA-: SPIRURIDA) IN AUSTRALIA 145 


culata From Verran, S. Aust; Arvarruty 

melinepy from Port Angusta, S$. Aust. 

Oregica gutiuralis from the Petermann 

Ranges, N.T. 

Although the hosts listed above cover a wide 
range of bird groups, and a wide geographical 
Tange, there appear to be no specific differences 
among the specimens from each. Although 
there is some variation in the position of the 
cervical papillac, and in the length of the cor- 
dons fn the male, there is often as much varia- 
tion betwoen specimens from one host as be- 
Iween specimens fiom different hosts. 


The cordons are short. They do not extend 
as far as the nerve fing in the male, or forther 
ihan the excretory pore in the female, 


The vulva is at about the mid-body, just in 
front of or just behind this, The vagina passes 
hackwards. Eggs are 38-39 by 21-23 jam, 

The caudal alae of the male are slender and 
only slightly wider in their anterior halves. 
There are typically four pairs and one median 
preanal papillae, seven pairs of  postanal 
papillae and a pair of very small phasmids, 
The postunal papillae are not atranged in two 
groups, hut lic progressively closer together 
towards the tip of the tail, In some specimens 
there are more or fewer papillae on one side 
or the other, bul these appear ta be ybnor- 
malities, The spicules are unequal; the tips of 
both are blunt and rounded. The species 
appears to be very close to 4. orieli Chabaud 
& Petter (1961), based on specimens from an 
vniole from Dahomey, which had been placed 
(with reserve) by Gendre (1912) in his species 
A. gracilis, from Recharge otra from the same 
locality. Gendre stules that the specimens from 
the onole were in nearly all points similar to 
those from the drongo, distinguished only by 
the number of postanal pupillae in the male, 
andl the shape of the tip of the male tail. The 
cordons of A. orioli are longer in both sexes, 
than those of the Australian specimens. In the 
absence of more information about A. orioli, 
the Avstralian specimens are regarded as a dis- 
linct species, In some ways it resenibles A, 
skrjebini tut differs from this species in the 
more slender build of the spicules, the un- 
enlarged tip to the right spicule, the shape of 
the caudal alae, and the detailed structure of 
the cordons, 


Acuaria colluricinclae n. sp. 
FIGS. 14-16 


Hest and locality: Coluricincla rifivensris 
from Eyre Peninsula, S. Aust, 


The material consists only of one male art 
one fomale specimen, but these differ distinctly 
from A. pelteras which appears to be the com- 
menest species of the genus jin AUstealian 
passerines. Mcasurements are given in Tuble 2. 

The cordons extend nearly to the posterior 
end of the glandular oesophagus in both sexes, 
at dittle hearer in the female. Detail of the cor- 
don structure ure shown in Fig, 14. 

The spicules are unequal in length; the right 
spicule ends in a swollen tip. There are four 
pairs and one median preanal papillae, 6 pairs 
of postanal papillae, and one pair of phasmids. 
The postanal papillae are asymmetrical { Fig. 
15), presumably an abnormal condition; the 
first 3 pairs are well spaced and spread over the 
anterior 220 ym of the 280 ~m long tail, while 
the Jast 3 pairs are Smaller and Jie on the ter- 
minal 50 jum, 


The species is distinguished from other Aus- 
tralian ones. by the ratio of the spicules, the 
structure of the right spicule, the grouping of 
the postanal papillae, and the cordon length. 
It is distinguished From other close species us 
shown in the key to species. 


Acuaria microecae n, sp. 
FIGS. 17-20 


Host and locality: Microeca leucophaea from 
Waikerie, S, Aust. 
The measurements of this species. of which 
onty [ male and | female are present, are given 
in Table 2. 


The cordotis of the male reach to the end of 
ihe muscular oesophagus; those of the female 
to about halfway between the head and the 
posterior end of the slandular oesophagus. 


There are four pairs and one medign pre- 
anal papillae, scven pairs of pustanal pupillae 
and a pair of subterminal phasmids. The post- 
anal papillae on each side are inure or less 
evenly spaced along the tail, the posterior anes 
slightly closer (ogether. The spicules are oqual 
in length and similar in build; each has 9 pair 
of short alae towards the distal end, and the 
rounded tip is bent ventrally, 

The vulva is slightly behind the midbody; the 
eggs gre 35 x 27 ym. 

The species is distinguished [rom others 
from Australia by the presence of equal spi- 
cules, It differs from other species in which the 
spicules are equal and in which there are 7 
pairs of postanal papillae, in having longer 
cerdons and in the very short spicules 


146 PATRICIA M. MAWSON 


um 
— 


12) 
oO 


B m 


100 


Figs. 14-16. Acuaria colluricinclae. Fig. 14.—Part of a cordon. Fig, 15.—Posterior end of male. Fig. 
16.—Tail of female. 

Figs. 17-20. A. microecae. Fig. 17.—Anterior end of male. Fig..18—Part of a cordon. Fig. 19.— 
Posterior end of male. Fig- 20.—Tail of female. 

Figs, 21-23. A. mirafrae. Fig. 21..-Anterior end of male, Fig. 22.—Parl of a cordon. Fig. 23. — 
Posterior end of male. 

Figs. 15, 19, and 21 to scale beside 21; figs. 16 and 20 to scale beside 20; figs. 17 and 23 to scale 
beside 23: figs, 18 and 22 to scule beside 22. 


ACUARTIA (NEMATONA: SPIRUREDA) IN AUSTRALIA 


Acuasia mirafrae n, <p. 
FIGS. 21-23 


Host and locality; Mirafrw javanica frora the 
Northern Territory. 


This collection comprises only two whole 
and one hroken male worms. Measurements 
are given in Table 2. 


The cordons extend a short distance behind 
the excretory pore. ‘The detail of the cordon 
pattem (Fiz. 22) w somewhat similar to that 
of A. petterue. 


There ate four pairs and one median preanal 
papillae, six pairs of postanal papillae and a 
pur of subterminal phasmids, The postanal 
papillae are arranged in two groups of three 
pairs. The right spreule ends bluntly and the 
tip is slightly indented, 


The species is distinguished trom A. peterde 
by the number of postarnal papillae and by the 
rather longer cordons, It is close to A. vractlis 


ww 


Genudre tm the body measorements, but ujlfurs 
in the spicule ratio and the arrangement of the 
pustanal papillae 


Acknowledgements 

Many of the birds dissected were given by 
the South Australian Museum or by the 
Northern Territory Muscum, [ am most grate- 
ful to the officers. of these: museums, Who look 
a great deal of trouble to get the bodies to. me. 
The Tasmanian ravens Were sent by Mr. Barry 
Munday of the Mi. Pleasant Laboratories of 
the Tasmanian Department of Agriculture, 
Luunceston. Specimens from exotic finches 
came from Dr. D. M, Murray of the C.S.1.R.0. 
McMaster Laboratory. Some hosts were cal- 
lected for me by colleagues—Mrs. Joan Paton 
and Dr, Michael Smyth. The single worm from 
Cervus sp. from Pearson |, was obtalacd in 
1933 by the Wood-Jones Expedition, T am 
gteatly indebted to all these helpers. 

The work was done in part under a Ruril 
Credits Development Grant. 


References 


Bovp, BE, M. (1956)—Two new species of 
stomach worms (Nematoda: Spirureidea) 
from the Blue Jay, Cyanecitta evistain L 
Proc. Helm, Soc. Wash. 23, 70-74. 

CHataup, A, G,, & Parres, A. (1961)~—-Néma- 
todes di genre AcHaria de la faune de France. 
Annly. Parasit. him. comp. 36, 409-424, 

Dusakoww, KF. (1845).—"Histoire ualurelle des 
Helminthes qu vers intesinaux”, [Paris] 

Erkurov, K, (1969),.—[A new nemialade, Aeiearin 
eremopiila (Sptruratu: Acuuritdac) from the 
shore lark] Mater. naveh, Knof. owes. 
Ohsheh, Gel’'mint, Year 1969, Pt, I, 339-343. 
(In Russian.) 

Genpar. EB. (1912).—Sur quelques espices de Dis- 
phirages du Dahomey. Proces-Verigua Sac. 
Linnéenne de Bordeaux 61, 23-31. 

Hsu Wet-Nnan (1963).—Studics on some parasitic 
fheniatodes of the order Spiruridu Chitwood. 
Act Zuel. Sinicg U5, 544-552. 

Jounston, T. H.. & Mawson, PM, (1941)}— 
Same nematode parasites of Ausiralian birds, 
Pree. Linn, Soe. NSW, 66, 250-256, 

Lent, H., Freitas, J, F. T,, & Proewes, M. C. 
£1945).—Algnns helmintos de aves colecio- 
nadas no Paraguai. Mem. Janse, Ovvalde Cruz. 
43, 273-285, 

Le Roux, P, L, (1930)—An Acinaria tAcvarta 
nartinagliai sp. nov.) fram a South African 
Weaver (Hyphantornis sp.). 16th Report 
Direct. Veter. Serv. Animal Indust. Union 
South Africa, 211-215. 

LENsTow, O. yon (1878)—Neuc Beobachtungen 
an Helminthen, Arch. Naturg. 44, 218-245. 

Mafcestone, P. A. (1931).—Parasitic nematodes 
obtained from animals dying in the Calcutta 
Zoological Gardens. Pts, 4-8. Rec.. Ine, Mus. 
33, 71-171. 

MAPLESTONK, P. A, (1931).—Parasilic nematodes 
obtained from animals dying in the Cajentta 


Zoological Gardens, Pts. 9-11. Ree. dra, Mire 
34, 229-261. 

Morerer, A. (1897)—Helminthelogische Mil- 
telungen. Arch, Naterg. 68, 1-26, 

Ozerska, Vo N, (1926).—Zur fauny Wer poru- 
siisthen  Wirmer der Wauspertinge des 
Dongehiets, Trudy Gor. Inst. eksp, Ver, 2, 
102-108. ; 

RASHEED, S. (1961)—The nematode parasites af 
the birds of Hyderabad (India), Biologie, 
Lahore 6, |-116, 

Rubpotert, C. A. (£819).—"Entozooruit synopnis 
cul accedunt. mantissa duplex el tnilices loco- 
pletissimi,'! [Rerolini.| 

Sench, SN. (1948),—Stuiies on the helminth 
parusites of birds in Hyderabad ‘State. Nem 
toda M7, J. Helminth 22, 199-218, 

SER TABIN, K. 1, Surkuogacaya, N. P., & Soporny, 
A. A, (1949) —{Spirtwata and Filariaty.] 
Opredilitel paraziticheshikh neniatad. Vol, 7. 
(Moscow. Akad. Sci. U.S.5.R.) Gin Russian, ) 

Searanin, K. LL, Soporev, A. A. & TVASHKIN. 
Vv. M, (1965),—[*Principles of nematology. 
Vol, 14, Sprrurata of animals and man und 
the diseases caused by them. Pr, 3, 
Acuarjidae."] (Moscow Acad. Sci. U.S.5.R,) 
(in Russian.) 

SKRIABIN. K. 1, Sonotev, A. A,. & TvAsHKIN, 
V. M. ¢1967),—["“Principles of nematology," 
Vol. 19, Spirurata. Pt, § (addendum),] (Mos- 
cow Acad. Sci. U.S.S.R.) (In Russian,) 

Wana, P. C. (1965)—Notes on Acuarioidea of 
birds from Fukien. China, Acta parasitol. 
S\nica 3, 15-29, 

Wititams, OL, (1929).—A crucial analysis of 
the specific characters of the genus Acuaria, 
hemiutodes of birds, with descriptions of new 
American species, Univ, Calif, Publs. Zool. 
33. 69-107, 


A LATE PALAEOZOIC GLACIATED GRANITE SURFACE AT PORT 
ELLIOT, SOUTH AUSTRALIA 


BY A. R. MILNES AND R. P. BOURMAN 


Summary 


A glaciated granite pavement, discovered at Port Elliot, South Australia, shows features that 
indicate an east to west movement of Late Palaeozoic ice, and thus significantly extends the known 
area of approximate east to west ice movement on Fleurieu Peninsula. The granite pavement and the 
profusion of granite erratics, together with the probable existence of an originally extensive pluton 
of Encounter Bay Granites, suggest the possible dissection of the pluton during the glaciation to 
form the present outcrop distribution. 


A LATE PALAEOZOIC GLACIATED GRANITE SURFACE AT PORT ELLIOT, 
SOUTH AUSTRALIA 


by A. R. Minnes* and R. P. BourMant 


Sunimary 


A glaciated granite pavement, discovered at Port Elliot, South Australia, shows features that indi- 
eale an eust to west moyement of Late Palaeozoic ice, and thus significantly extends ihe known area 
of approximate cast to west ice movement on Fleuriet. Peninsula, The granite pavement and the pro- 
fusion of granite erratica, together with the probable existence of an originally extensive pluton of 
Encounter Bay Granites, suggest the possible dissection of the pluton during the glaciation to form 


the present eutcrop distribution, 


futroduction 


Fleurieu Peninsula, South Australia, has for 
many years been studied from the point of 
view Of iis sculttered exposures of glacigene 
sediments (now known to extend over a large 
proportion of South Australia} and its Jand- 
forms, many of which have been accorded a 
elacial origin. Such investigations (e.g, How- 
chin 1898a, 1898b, 1908, 1910a, 1910b, 1926, 
1929; Campana & Wilson 1955) were initiated 
by the discoverics of placiated Cambrian and 
Proterozoic bedrock surfaces in the Inman 
Valley (Selwyn 1859) and at Hallett Cove 
{Tate 1889). Crowell & Frakes (1971) sum- 
marised and in part reinterpreted many of the 
earlier investigations in the context of a 
regional study of the Late Palacozvic glaciation 
in Australia. 


The age of the glacigene sediments in many 
localities in South Australia was determined 
by Ludbrook (1956, 1967, 1969:2,b) trom fora- 
miniferal, studies ay lower Permian, Tentative 
evidence for a Late Carboniferous to Lower 
Permian age for the glaciation, in the form of 
a possible Late Carboniferous microflora in gla- 
eigene seuitnents in the Lake Phillipson bore in 
north-west. South Australia, was. recorded by 
Balme (1957). Harris & McGowran (1971) 


recently described palynomorph assemblages 
of Permian age in glacigene sediments from 
several localities adjacent to Fleurieu Penin- 
sula, and in addition, recorded a reworked 
Devonian microflora in glacigene sediments 
from Waterloo Bay on Yorke Peninsula, For 
the purpose of the present paper, we will refer 
to the age of the glaciation as Late Palaeozoic. 

Our contribution was initiated by the dis- 
covery of a glaciated granite surface at Port 
Ethot, and other glacial features throughout 
Fleurieu Peninsula, These finds have provided 
significant information on the local effects of 
the Late Palaeozoic glaciation, and the direc- 
tion of movement of the ice. 


The Encounter Bay Granites 

The Encounter Bay Granites! ccop out in 
the Encounter Bay area as discontinuous 
masses along the coasiline, and on the adja- 
cant small islands, The contact of the granites 
with the surrounding Kanmantoo Group meta- 
sedimentary recks is exposed in only. two loca- 
lities, Rosetta Head and Wright. Island, In all 
other localities, it is obscured cither by Late 
Palaeozoic to Recent sediments, or by the sea. 
Several Jarge xenoliths of unaltered Kanman- 
too Group metasedimentary rocks within the 
granites: are possibly roof pendants, and may 


* CSIRO, Diyision of Soils; Glen Osmond, 8. Aust, 5064. Formerly at; Department of Geology and 

Mincralogy, The University of Adelaide, 8. Aust. 

1 Department. of Geography, The University of Adelaide, Adelaide, S. Aust, 5000. 

IEncounter Bay Granites is the term used to describe collectively the several tclated varieties of 
granitic rocks that crop out in the Encounter Bay area and al Cupe Willoughby, Kangaroo Island. 
It is intended that it replace the synonymous term “Eyjcanater Ray Granite” defined by Dasch, 
Milnes & Nesbitt (1971). 


Trans, R. Soc. S, Aust. Vol. 96, Part 3, 31 August 1972- 


150 A, R. MILNES AND R. P. BOURMAN 


STRIAE DIRECTION OF LATE PALAEQZOIC 
GLACIATION-FLEURIEU PENINSULA 


N 


, THOR 
“Es DIRECTION OF ICE 


MODYEMENT 
5 0 5 ia Kms 
——— 


f ‘FLEURIEU 
yy, PENINSULA 


ST 


ia Fale 
TWokaverta Ck. 
<—-S 
Gurremy Ck, 


C ELLOT 


Lyicton HARBOUD 
SEGA AN ITE LS. 


Awaishr ha 
Fe ROSETTA HEAD 
CKING BEACH 


A} 
(y WESI 15. 


/ ENGOUNTER 
fy 
\ . ~ BAY 
CAPE eee ee andl 
JERVIS “SS or 


Fig. b 


Map showing the known occurrences of Permo-Carboniferous glacialed pavements and cor- 


responding striae directions, Fleurieu Peninsula, South Australia. 


indicate the close proximity of the present 
level of exposure to the roof of the original 
intrusion. Thus, the granites from West Island 
to Port Elliot appear to have originally formed 
part of the north-western wall and roof of a 
large pluton, which may have extended for 
some distance eastwards in the present position 
of the Southern Ocean and the western portion 
of the Murray Basin. 

The granite outcrops show considerable 
modification as a result of surface weathering 
and erosion. Features such as sheeting, flared 
slopes, and gnammas, all of which are chatae- 


teristic of inselbery structures, have been ob- 
served in many localities, In addition, a marked 
development of tafoni, micropediments. and 
corestones has been noted, 


The Glacial Pavement and Associated Features 
at Port Elliot 

An undulating smoothed and polished gra- 
nite surface occurs at the edges of a pathwuy 
excavated across the top of the promontory 
between Knights Beach and Green Bay, Port 
Elliot (Figs. 1-3). The surface is poorly ex- 
posed (Figs. 4, 5) but crops out intermittently 


PALAEOZOIC GLACIATED GRANITE 


Low level oblique aerial photograph of 
the promontory between Knights Beach 
and Green Bay, Port Elliot, on which the 
glaciated granite pavement is exposed. 


Fig, 2. 


~PORT ELLIOT 


‘pavement 


Pavements 


glacigene silts xenalith 


granite 


Fig. 3. Geological sketch of the same locality as 
shown in Fig. 2. 


151 


over a distance of approximately 30 m, and 
is extremely variable in orientation. It is 
overlain by thinly to very thinly bedded silts. 

The area of exposure, corresponding to the 
easternmost extent of the pavement, is less than 
| m square. However, its extension for at least 
| m further westwards has been proved by 
minor excavation, Well preserved striations and 
grooves and possible crescent-shaped gouges 
were observed on the pavement in this ex po- 
sure only (Fig. 4), and indicate a direction of 
movement of the glacial ice from approxi- 
mately east to west (255° to 260°). These sur- 
face features have been partly obliterated 
(probably by traffic along the pathway) on 
those parts of the pavement exposed for 
some lengih of time, but are extremely well 
preserved on that part of the pavement recently 
exhumed from beneath a cover of silts. 

The sediments overlying the pavement are 
well exposed in the cutting at the edge of the 
pathway across the top of the promontory, and 
are similar to sediments described by earlier 
workers as glacigene from other parts of 
Fleurieu Peninsula. They consist of thinly to 
very thinly bedded brown to red and greyish 
white coloured silts with minor grit bands and 
lenses. Many beds show extremely fine lamina- 
tions. Bedding attitudes vary from horizontal 
to a shallow dip towards the north-west, but in 
view of the variability of the underlying sur- 
face, are interpreted as primary depositional 
attitudes, 

Immediately beneath the granite pavement, 
there is a weathered zone of varying thickness 
characterised by the presence of innumerable 
small-scale fractures which appear to have 
mainly developed parallel to the pavement sur- 
face (Fig. 5). In addition, a skin of goethite 
up to I cm thick has formed on parts of the 
pavement (Fig. 6). These features are a result 
of recent weathering. 


The glacigene silts are directly overlain by a 
calcareous conglomerate, up to 1 m_ thick, 
which infills fractures in the uppermost beds of 
the silts (Figs. 7, 8). The conglomerate is in 
turn overlain by a consolidated calcareous 
beach sandstone which crops out spectacularly 
in the cliffs at the back of Knights Beach, The 
sandstone is capped by a layer of massive to 
nodular calcrete, and this laps directly on to the 
granite on the western and eastern margins of 
the promontory (Figs. 2, 3). 

The pavement and overlying glacigene silts 
are presently about 10 m above sea level. How- 
ever, it is clear that they have been at or below 


A. R. MILNES AND R, P, BOURMAN 


et 
wy 


PALAEOZOIC GLACIATED GRANITE 


sca level becuuse of the nature of the over. 
vine sediments. “(he conglomerate is jinter- 
preted as the resull of reworking of both the 
placizene sediments and the beach sandstone, 
por to ihe formution of the calcrete crust. 
Furthermore, it seems likely that the reworking 
occurred after the main period of heach deve- 
lopment, but in a similar although somewhat 
more sheltered environment, pethaps protected 
trom the lull crosive capacity of ihe sen hy the 
outcropping granite. In view of the thick deve- 
lopment of sandstone immediately inland from 
the granite, it is probable that a considerable 
proportion of the glucigene sediments ori- 
ginally present in this focalily has been re- 
moved. 

The pavement apparently defines part af the 
southern murgin of 9 focal glacigene sediment- 
filled busin, which occupies the area between 
the stanite outcrops at Port Elliot and the hills 
of Kanmantoo Group metasedimentary rocks 
3 km cto the north. Glacigene sediments have 
been logeed to depths of 175 m and 91 m in 
twa bares drilled in this basin to the north- 
cast of Port Elliot (Crawford & Thomson 
1959), However, the sediments do not crop 
ouc.at the surface as mapped on the Encounter 
I-mile sheet. being cflcctively blanketed by 
fore recent deposits including outwash, allu- 
vium and, nearer the coastline, consolidated 
(7) Pleistocene beach-dune sandstones and sine 
consolidated presemt-day beach and dunce sands. 


Other Glacigenc Sediments in the Port Elliot 
District 

Further outcrops of glacigene sediments, 
consisting of yery thinly laminated white tu 
buff coloured clays and associated sunds with 
sporadic ertatics, occitr in a Narrow north 
easterly trending valley just over 4 kim north of 
Port Elliot township. The erratics include 


“Brock, E. J. €1964)—The denudation chronology 
(partly unpublished). 


153 


exotic shanites, gneisses and quartzites, to- 
ether with boulders of locally derived meta- 
sandstone, but there is no evidence of erratics 
ef the nearby Encounter Buy Granites, One 
boulder of coarse grained gneiss contains bive 
Opalescent quartz, and is similar to rock-lypes 
found jn the alder Proterozoic basement inliers 
north-west of this locality, 


Discussion 

The glaciated granite pavement at Port Elliot 
is significant with regard to an interpretation 
of the Late Palseozoic to Recent geological and 
ecomorphological history of the Encounter Bay 
area, especially in relation to the nature and 
distribution of the outcrops of Encounter Bay 
Granitcs. The pavement Sinks with the glaciated 
Kunmantoo Group metasedimeitary fack 
pavements throughout Fleuricu’ Peninsula in 
Tevealing glimpses of u recently exhumed Late 
Palaeozoic landscape. Furthermore, tt indicates 
thut the Encounter Bay Granites were exposed 
at the carth’s. surface during and possibly prior 
to the glaciation, and ntar the same level 
within the intrusion us presently exposed, Adili- 
tional evidence for their exposure ix provided 
by the profusion throughout Fleurieu Penin- 
sula of erratics of Encounter Bay Granites, 
which are distinguished by their characteristic 
opalescent blue quartz crystals, 

The direction of ice movement determinvd 
from the aurface features preserved on part of 
the pavement is consistent with the dircctions 
determined in a similar fashion from 12. ghu- 
ciated Kanmantoo Group metasedimentary 
Tock surfaces (Fig. 1) on a wide yariety of 
topographical features (Howchin 1926; Brock 
19642, Maud 19678, and unpublished field 
observations). and thus we are able ta extend 
the known urea Of approximate east to west 
movement of the Late Palacozoic ice. At 


of Fleurieu Peninsula. M.A, thesis Univ. Adelaide 


8Maud, R, R. (1967}—The Permian of the arex ardund Mount Compass, (Unpublished maniécripl, 


CS.LR.O, Division af Soils, Adeluide.) 


granite pavement, showing parallel! striae, 


Steeply dipping portion of the granite pavement (Pv) exposed edge-on near the ceptre of 


photograph. Zone of disintegrating granite (Dg) between the pavement and fresh eramile 


Contact between glaciated granite pavement and overlying glacigene silts (Gs). Thin skin of 


goethite (Go) covers the pavement surface al the base of silts. Disiniegrating sranite (De) 


Fig, 4, The eastern-most exposure of the glaciated 
Fig. 3. 
(Fg). Scale represents 20 cm, 
Fig. 6. 
7 beneath pavement. Scale represents 10 cm. 
ig. 7. 
Tepresenis 10 em. 
Fig. &. 


Calcureaus conglomerate (Ce) infilling fractures in upper beds of glacigene silts (Gs). Scale 


Section showing glacigene slis (Gs} at base (dipping al shallaw angle to the west), overlain 


by calcarcons conglomerate (Cc) amd a massive calcrete (Ca) crust. Scale represents 30 em, 


134 


Penneshaw (Chrisumas Cove) on Kangaroo 
Island, Ward (1922) recorded a restricted ex- 
posure of glacigeng sédiments and a smoothed 
and striated Kanmantoo Group  metasedi- 
mentary rock surface that also indicated an 
“eust-west cirection” of ice movement, ‘This 
Hirectlon, however, is at variance with that 
measured at Hallett Cove (Howchin 19246; 
Crowell & Frakes 1971) on one of the two 
other glaciated pavements in South Australia 
for which there is published infornration. Here 
the indicted direction of ice Movement is to- 
wards the north-north-west. Pritchard (1892) 
describes! elicited surfaces with similar north- 
south oriented striae on Cambrian limestone 
wear = Corramulks on Yorke Peninsula. 
Althuugh there are slacigene sediments in the 
area (Crawford £960), the origin of the typical 
Hat limestone outcrops has been. misinterpreled 
by Pritchard. 


Strive directiuns un vhe Huallew Cove pave- 
ments are jinomulois when compared with 
those measured from the numerous pavements 
In the south. Moreover, Hallett Cove is sepa- 
rated from the pavement localities of Fleuricu 
Peniosula by at least two mayor Faults, along 
which there has been marked vertical displace- 
ment during the Tertiary (Thomson & Horwitz 
1962), Providing there has been no pust- 
alacial rotation of the Proterozoic rocks, the 
swe Ulrections and the occurrence of erraties 
of Encounter Bay Granites. st Hallett Cove 
nay Nave resulted from either: 


(a) Che suuth to north mouvement of an ini- 
tial vontinental ice sheet over Fleuricu 
Peninsula and its immediate environ- 
men! as suggested hy Crowell & [rakes 
11971), followed by the later develop- 
ent of w Westerly moving ice Sheet; or 


{b) the effect of topography on a noorth- 
westerly moving ice sheet. 


The first possibility requires that north-trend- 
ing striae initially developed on bedrock sur- 
fnces on Fleurieu Peninsula were obliterated 
in all localities but Hallett Cove by younger 
westerly moving ice. Ip thus introduces the con- 
copt of multiple glaciation, but there is as yet 
ne unequivocal evidence for this in South Aus- 
Irwha. Neither the crossing striae Observed on 
many pavements, nor the striated pavement 
developed on glacigene sandstonc near the 
bank at the Finniss River (Fig. 1) require 
multiple glaciation (Flint 19571, The remark- 
alle consistency of striae directions on Fleurieu 
Peninsula and sastern Kangaroo Island over 


A. R. MILNES ANT R. Po BOURMAN 


varied lopography does not support the coo- 
tept of u system of irregular valley glaciers 
proposed by Campuna & Wilson (1959) and 
Crowell & Frakes (1971), hut the movement 
of a thick nil extensive ice shect. 


Evidence for the direction of ice amuvement. 
it addition tu strise which are usually only 
reliable indicators of local ice movement, is 
provided by the chsiributiun of certain ertatics. 
In particular, thé occurrence in glacigene sedi- 
ments on Fleuricu Peninsula of uw distinctive 
feldspar-quanz porphyry and af coarse grained 
non-folinled ted granites, apparently derived 
from the suite of granitic rocks that crops out 
between Murray Bridge and Dergholm {south- 
western Victoria), indicates un overall north- 
westerly icc Movement, af we assume that the 
present oulcrop distribution of these granites 
approximiutes to their real extent. ‘Vhis direc- 
tion is Close io the averaye of the Fleurieu 
Peninsula and the Huyllett Cove striae direc- 
tans, 


Thus, although the striate un Fleurieu Penin- 
sula pavements are constant in dircetion over 
considerable topographic relief, lopupgriphy of 
a much greater maynitude during the Late 
Palacozore could have channelled  nurth- 
Werterly inoving ice in a westerly direction 
across Pleurice Peninsula, The possihility of 
deflection of the ice by a north-easterly trend- 
ing vulley at Hallett Cave has lonz been con- 
sidered (Sprigg 1942). Such topographic in- 
fluences on a north-westerly trending ice sheet 
seem to account for the observed facts. 


We inierpret the present distribution of the 
outcrops of Encounter Bay Granites as the 
result of dissection of an originally extensive 
pluton by the exploitation, either of structural 
features such a jointing or of pre-existing 
druinuge valleys, hy westerly moving Lute 
Palasozaic tee. Evidence for glacial erosicn 
includes. che pavement ul Port Flliet and the 
abundance of granite erratics within the glaci- 
gene sediments throughout Fleuricu Peninsuli, 
In fact, the granite oulcrups seem ta have been 
preserved until recently beneath a cover of 
g@lucigene sediments, the reworking ul which 
has prociicecdl the conspicuous boulder fields 
acen for example near Rosetta Head and King 
Beach. The occurrence of many granite lypes 
foreign tothe Encounter Bay Granites in these 
ficlds docs not support the suggestion by 
Crowell & Frakes ¢1971) that the boulders in 
these localities were derived from nearby out- 
crops by mass-wasting or by the action of 
SLOT Wives, 


PALAEOZOIC GLACIATED GRANITE 182 


Conctusions 


The smoothed ati striated granite surface 
at Port Elliot, together with 12 glaciated Kan- 
mantoo Group metasedimentary rock surfaces 
throughout Fleurieu Peninsyla and one on 
Kangaroo Island, exhibit a remarkable consis- 
Iency of striae directions oyer a variety of 
topographic forms. This can only be the effect 
of a thick ice sheet, which is shown to have 
moved westwards over Fleurieu Peninsula. The 
striae directions measured from glaciated sur- 
faces at Hallett Cave are anomalous in this 


regard. However, these differences may reflect 
lopographic influences, 

The present broad distribution of outcrops 
of Encounter Bay Granites is interpreted pri- 
marily as the result of ice action. although 
there is abundant evidence of considerahle later 
modification of some outcrops.. 


Acknowledgements 
We Wish to acknowledge the helpful criti- 
cism of Mr. G. Blackburn, and the comments 
ot Dr. V. Gostin and Dr. B. Daily ou an early 
draft of the paper, 


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ADDITIONS TO THE HYLID FROG FAUNA OF NEW GUINEA, WITH 
DESCRIPTION OF A NEW SPECIES, LITORIA TIMIDA 


BY M. J. TYLER AND F. PARKER 


Summary 


Litoria timida, a new species of New Guinean hylid frog of the Litoria dorsalis group, is described 
and reported from five localities, ranging from the headwaters of the Fly River to the vicinity of 
Port Moresby. Three additional species of Litoria, formerly known solely from Australia, are 
reported from localities adjacent to Torres Strait. The diagnostic characters of each species are 
described, together with notes on habitat preferences and habit. 


ADDITIONS TO THE HYLID FROG FAUNA OF NEW GUINEA, 
WITH DESCRIPTION OF A NEW SPECIES, LITORIA TIMIDA 


by M. J. Tycter*® and F. PARKER? 


Summary 


Litoria timida, a new species of New Guinean hylid frog of the Litoria darsaliy group, is described 
and reported from five localities, ranging from the headwaters of the Fly River to the vicinity of 
Port Moresby, Three additional species of Litoria, formerly known solely from Australia, are reported 
from localities adjacent to Torres Strait. The diagnostic characters of each species are described, 
logether with nutes on habitat preferences and habit. 


Introduction 


The portion of the island of New Guinea 
adjacent to Australia, now within the Western 
District of Papua, is.of considerable zoogeo- 
graphic interest because of the close faunal 
similarity with the Cape York Péninsula. of 
Queensland. Although the first publication on 
a collection of frogs from this part of New 
Guinea appeared nearly a century uyo 
(Macleay 1878), this and sabsequent contribu- 
tions (Roux 1920; Loveridge 1956) dealt with 
very small samples and, until recently, the 
extent of the frog faunal similarity to Queens- 
land remained uncertain (Tyler, in press}. 

During three years residence at Daru one 
of us (F.P.) has collected several thousand 
frogs throughout the Western District. Here we 
confine our attention to some of the representa- 
tives of the Hylidae in this collection, reporting 
details of the distributions and habitats of three 
species formerly known to occur only in Aus- 
tralia, and describing one new species, Details 
of the field expeditions that resulted in the 
collection of same of the specimens have been 
published elsewhere (Parker 1970). 


Methods 

The specimens forming the subject of the 
paper are deposited in the collections of institu- 
tions ahbreviated in the text as lollows: 
American Muscum of Natural History 
(A.M.N.H.); Australian Museum (A.M.); 
Museum of Comparative Zoology (M:C.Z.): 
Papua and New Guines Museum (P.N.G.M.); 
South Australian Museum (8.A.M,); Depart- 
ment of Biology, University of Papua and New 


Guinea (U.P.N.G.). The letters FP. in paren- 
theses preceding specimen teference numbers 
indicate that these are field numbers for cur- 
rently unregistered material, 

Methods of measurement and morphological 
and geographical descriptive terminology fol- 
low those of Tyler (1968a) for species then 
referred to the cosmopolitan genus Ayla. Such 
species from New Guinea and Australia com- 
prise an endemic genus fer which the name 
Lileria has heen proposed (Tyler 1971), The 
descriptive abbreviations used are E-N (dis- 
tance between the eye and the nwris)) IN (ine 
ternarial span); HL, (head length); MW (head 
width); S-V (snout to vent length); TL. (tibia 
length). 

Western District localities cited in the text 
are shown in Figure L. 


Litoria timida n. sp. 

Holatype: S.A.M. 11658. Am adult: mule 
collected at Menemsorae, Western Dis- 
trict, Papua New Guinea by F, Parker on 
30 March, 1969. 

Definition: An extremely small lowland species 
{males 21.3-23.9 mm; females 26.3 mnv) 
characterised by its elongated head, curved and 
extremely sharply defined canthus rostralis, 
farge and prominent eyes, unwebbed. fingers 
and reduced webbing between the tocs, 

Descriplion of Holetype: The head is flattened, 
angular and distinctly longer than broad 
(HL/HW 1,246), its length equivalent to more 
than one-third of the snout to vent length. The 
snout is extremely prominent. and angular when 
viewed from above, prominent and projecting 


* South Australian Museum, North Terrace, Adclaide, S. Aust. SO0G- 
+ P.O, Box $2. Daru, Wesicen District, Papua New Guinca. 


Trans. R. Soc. S. Aust. Vol. 96, Part 3, 31 August 1972, 


158 M, |. TYLER AND F. PARKER 


Derongo) ® 


Yongtau ({ @ f 
Ningerum 


Mendua 


Kiunga 


L 
*Wearh % 


4 (MBre ox 


50 km 


| 


Tig. 1. 


far beyond the anterior limit of the mandible 
in profile. The nostrils are dorso-lateral, their 
distance from the end of the snout being con- 
siderably less than that from the eye. The dis- 
tance between the eye and the naris is yery 
much greater than the intermarial span (B-N/ 
IN 1,527). The canthus rostralis in long, well 
defined and curved, and the loreal region 
steeply sloping. The eye is very large and 
prominent, its diameter slightly less than the 
distance between cye and naris. The tympanum 
is prominent, having a diameter equivalent to 
approximately one-half of the eye diameter, 
and sepurated from the eye by a distance 
cquivalent to one-half of the diameter of the 
tympanum. The vomerine teeth ure on two 
prominent, raised and slightly oblique scries 
between the choanac, The tongue is small and 
circular. 


The fingers ure Jong, slender, unwebbed and 
luck literal fringes, in decreasing order of 
length 3>4>2>1. The terminal discs are 
large, the diameter of the disc of the third 
finger being twicc the diameter of the pen- 
ultimate phalanx. 


2 7 ROld Zim 
Wipim_e Kuru 


Collecting sites in the Western Distrizt, Papua. 


The hind limbs are long and slender with a 
TL/S-V ratio of 0.605, and toes in decreasing 
order of length 4>5 — 3>2>1. The webbing 
between the toes is reduced, reaching the sub- 
articular tubercle at the base of the penultimate 
philanx on the third and on the fifth toe, There 
is a prominent, elongate imner metatarsal 
tubercle and a prominent raund ouler meta- 
tarsal tubercle. 


The dorsal surfaces of the head, body and 
limbs bear scattered and poorly developed 
tubercles. The throat and chest ure smooth, 
and the abdomen and posterior ventral sur- 
faces of the femora granulur. 


This male specimen has a single, subman- 
dibular vocal sac and glandular, but unpig- 
mented, nuptial pads. 


The dorsum is dull brown and mottled with 
slightly darker and irregular patches. ‘The 
ventral surfaces are very pale cream with a 
brown reticulum on the throat and chest. There 
is fine light stippling on the ventral und pos- 
terior surfaces of the thighs, Dimensions: 
Snout to vent length 21.5 mm; tibia length 


HYLID FROG FAUNA OF NEW GUINEA 159 


Fig. 2. 


13 mm; head length 8.6 mm; head width 6.9 
mm; eye to naris distance 2.9 mm; internarial 
span 1.9 mm; eye diameter 2.7 mm; tympanum 
diameter 1.5 mm. 


Variation: The paratype series consists of 
sixteen adult males and one gravid female: 
M.C.Z. 82380-90, S.A.M. 11659-61, collected 
at Menemsorae with the holotype M.C.Z. 
82400, Derongo; M.C.Z, 82377-78, Yongtau. 
The series forms a remarkably homogeneous 
group, the snout to vent length range of the 
sixteen males being only 21.3-23.9 m. The 
snout to vent length of the female is 26.3 mm. 
Similarly, there is scant variation in their pro- 
portions: TL/S-V 0.605-0.647 (mean 0.627), 
E-N/IN 1.381-1.722 (1.577), HL/HW 1.114- 
1.348 (1.245), HL/S-V 0.376-0.413 (0.399). 


In all specimens the canthus rostralis is con- 
sistently a prominent feature, the eyes are large 
and protruding and vomerine teeth are present. 


With two exceptions the paratypes are 
brown dorsally with or without faint and 
irregularly shaped paler markings. The excep- 
tions are grey; one uniformly so, whilst the 


Litoria timida, paratype (M.C.Z. 82390). 


other (M.C.Z. 82400) bears two very broad 
and clearly demarcated, pale, dorso-lateral 
stripes extending from the eye to the groin. 
Pigmentation of the ventral surface of the 
throat and body varies from a very light stip- 
pling to extensive dark brown reticulations on 
the throat, chest and anterior half of the 
abdomen. 


In life the dorsum is brown with paler mark- 
ings, and the posterior surfaces of the thighs 
are dark brown with or without a few yellow 
spots. The labial stripe is white, and the iris 
a reddish bronze, being palest around the pupil, 
The flanks vary from brown to pale yellowish, 
and the groin is bright yellow. The ventral sur- 
faces are yellow and the ventral markings re- 
ported above are brown. 


Ten additional specimens have been col- 
lected from localities to the east of the Western 
District: M.C.Z. 82379, Oroi, Purari River 
[Lat. 7°237S., Long. 145°11’E.], Gulf. of 
Papua (obtained by F.P.): S.A.M. 10656-8. 
11657; U.P.N.G, 2510-11, 2581-3, Brown 
River Forest Reserve, north of Port Moresby 
(collected by J. I. Menzies). They differ from 


160 M. J. TYLER AND F. PARKER 


he type series in the following respects: the 
posterior surfaces af the thighs and the groin 
bear a striking pattern of yellow markings on 
a dark brown background, and several exhibit 
a very narrow, light, mid-vertebral stripe, 


Comparisan with other species: Literia timida 
is @ member of the Literia dorsaliy group, de- 
fined by Tyler (1968a). which contains L. 
dorvaliy of New Guinea and L. rmierobelos ot 
northern Queenslind. The members of the 
group shure long, slender, unwebbed fingers, 
reduced interdigital webbing of the toes and 
extremely small adult size. 


Litoria timide is larger than L. dorsaliy and 
L. microbelos (maximum adult size of males 
and females: dersalis 21 mm and 22 mm; 
microbelos 194 mm and 18.9 mm), The 
feature by which these species can be ¢dis- 
linguished most readily is the canthus Tostralis: 
strongly curved and sharply defined in rina, 
but straight or slightly curved and not a 
prominent feature in L. dorsalix and L. niiero- 
belos (Fig. 3), From £. nicrobelos the new 
species is Further distinguished by its coloura- 
tions examples of L, micrabelos are usuidly 
only very slightly stippled and there is @ ten- 
dency for the dorsodateral surfaces ta be 
durker than the dorsal, In £. tinder plgmenta- 
tion is more extensive, and in the only speci- 
nen in which the dorso-lateral surfaces dilfer 
fran) the dorsal, the dorsal portion is darkest, 
Litera = mitrobelas lacks yomerine teeth 
(present in £, Unda) and has much shorter 
hind fimbs, as indicated by a comparison of 
the ranues of the TL/S-¥ ratios: 0.605-0.647 
in L. Moda, 0.500-0.579 in a series of thirteen 
1. micrahelos from Cainns—S.A.M_ 12571-4, 
M.C.Z. (Uneat.). 


| / ( UH 
A B 


Heads of A, Litoria timida and B, L. 
dorsalis, demonstrating the difference in 
ihe shape of ihe canthus rostralis. Both 
specimens are males and there is no evi- 
dence of sexual dimorphism. 


Literia’ dorsalix resembles L. pierahelas 
more closely than it does L. /imida, Pattern of 
pigment on the dorsum of L. dorsaliy most 
commonly tends to form broad and slightly 
contrasting, longitudinally vrranged. light and 
dark stripes. Womerine teeth are present but 
diffieult ta detecL when elevations of the 
vomerine banes are absent. The range of TL/ 
§-V ratios is intermediate belween those of 
L. timida and L. inierebelos (0.541-0,621). 


The only other hylid species recorded trom 
the Southern Lowlands of New Guinea 
(recognised as a faunal unit by Tyler (19683), 
and including the Western District of Papua) 
which approximate the size of L. rinida are 
1. hicolor and associated species. Such species 
are usually uniformly bright green dorsally. 
and can be further distinguished from L. tide 
by possession of webbing between the fingers. 
Jateral fringes on the fingers. and fully webhed 
toes 


The high TL/S-V raho ot L. tinida ips 
prouches that of Lo meyuta, Because the first 
couplet in the relevant key to hylid species 
compiled by Tyler (1968u) involves separation 
of species on the basis of 9 TL/S-V_ ratio 
higher than or lower than 0.65, comparison 
with L. nasura is mecessary. In habitus these 
Species are quite distinet, L. nasa being much 
larger and having o very long. slender head 
wd body Adult L. nasare attain 45 om 
(males) and 55 mm (females), juveniles (und 
adults) can be readily distinguished by their 
possession of longitudinal skin folds on the 
dorsum, 


Haliui: The type locality of Menemsorae and 
Derongo and Yongtau ute in the viemity of 
Ningerum at the headwaters of ihe bly River 
in the extreme northwestern part of the West- 
ern District (Pig 1), and are within dense rain 
forest. The lerrain is gently undalating to the 
south of Ningerum and hilly to the north. The 
series from Menemsorac were collected on the 
leaves of plants adjacent to, or overhanging, 
the water of a permanent swamp in the forest, 


Oroi is situated in tropical rainforest partly 
cleared to form garden areas, It is believed tht 
the specimen obtained there had been collected 
in the extensive adjacent saga swamps. The 
Brown River Forest Reserve similarly provides 
an extremely moist habitat. 


Call: Calling males were observed occupying a 
horivontal position on leaves close to the water. 
We lack tape recordings of the calls, but note 


RYLID FROG FAUNA OF NEW GUINEA 16] 


that audibly they are feadily distinguishable, 
The call of L. timida was likened to two short 
clicks, each followed by a pause, and then a 
sequence of tapid notes: “tik... . tik. 2.4, 
Lindiititi’. In contrast, the call of £. dorsalis 
comprises a series of from six to twenty short, 
“woodeny buzzes”, and that of ZL. microbeloy a 
low, imsect-like “creeeek”, becoming pro- 
gressively higher pitched. 


Notes: It was noted that indigenees of the 
Ningerum area did not recognise the new 
‘specics, They consistently considered specimens 
to be juveniles of larger species such as L. 
thesaurensis or L. amboinensis, 


The specific name proposed for the species 
alludes to the timid nature of the animal. 
When disturbed it rapidly attempts to escape- 


ADDITIONS TO THE PAPUAN HYLID 
FROG FAUNA 


Litoria nigrofrenata (Gunther) 


Until recently this species was known solcly 
from the types collected at an unspecified 
locality on the Cape York Peninsula, re- 
examined by Moore (1961) and Tyler 
(1968b), Additional specimens were sub- 
sequently collected by the second author on 
the Cape York Peninsula at Endeavour River, 


Leggits (Jones) Lagoon and Big Tableland (all 
within a 15 km radius of Cooktown). 


The occurrence of this species in New 
Guinea is based on the following nineteen 
specimens: 


Boze, Binaturi River: M.C.Z. 81097; Kuru: 
M.C.Z. 81094-96, P.N.G.M. (F.P.) 438 (2), 
§.A.M. 11408; Morehead: M.C.Z, 84549-50, 
S.A.M. 11316-19, 10660, [1467 (3}: Old Zim. 
Oriomo River: M,C.Z. 79723-24, S.A.M, 
10621 (2); Weam: M.C.Z. 79725-26. These 
localities are all situated to the south and south- 
west of the outlet of the Fly River and are 
within 60 km of the coast. 


The diagnostic characters of this species arc 
as follows: Size moderate {maximum snout to 
vent lengths; males 41 mm, females 42 mm). 
Head elongate with the snout projecting in 
profile; head consistently longer than broad. 
Fisst finger longer than second; fingers without 
webbing; fourth toe webbed to base of penulti- 
mate phalans; terminal discs small, Dorsal sur- 
Faces of the head, body and limbs uniform pale 
hrown, with a broad, dark brown stripe extend- 
ing from the tip of the snout to a position 
posterior to the iympanum. There is a hori- 
zontal white bar anterior to the cye partially 
bisceting this dark lateral band (Fig. 4). 


Because few detailed comparisons of such 
disjunct populations have been undertaken and 
none previously published, measurements and 
proportions of specimens of L. nigrofrenata 


Fig. 4, 


5mm 


Lateral view of head of Litoria nigrofrenata: 


1a M_ 1. TYLER AND F. PARKER 


Fram New Guinea and Australia are presented 
below. 


AUSTRALIA NEW GUINEA 
Number 12 5 
S-V 37,2441 36,9--38.8 
(nsales) (mean 38.2) (mean 38.0) 
SV 38.9- 41.3 414 
(females) (mean 40.1) (mean 41.8) 
Th /S-¥ 0,.541-0,697 0,584-0.666 
(pooled) (mean 0,629) (meun 0,632) 
FN/IN 0.925-1.162 0.975-1.083 
{pooled} (mean 0.999), (mean 1,007) 
HI. /HW 1.078-1.309 1481--1.262 
(pooled) (mean 1.180) {nyean 1.230) 
HL/s-V 0,360,398 387 0.404 
(pooled) (mean 0.376) {ime 4.396) 


In life the dorsal surfaces vary from a pale 
yellowish-brown ta reddish-brown; adults are 
immaculate, but juveniles are speckled wath 
darker pigment. 

Liroria nigrofrenaa exhibits considerable 
variation in leg length and individuals with 
TL/S-¥ ratios equal to or exceeding 0.65 key 
out in Tyler's (19684) key of lowland species 
lo L. nasutea. Specimens with lower TL/S-V 
ratios key out to L. vagabuneda. 


Lueria nasuta is similar in habitus to LZ. 
nivrofrenate but is distinguishable by possession 
of longitudinal skin folds on the dorsal surface 
af the hody and by lack of a lateral head 
stripe, In contrast, L. vegabanda is a broad- 


headed species with short limbs (TL/S-V of 


allolype — 0.513) (Tyler L968a), Literie werge- 
bunda is dark blue in preservative and so pro- 
hably green in life, 


Litevia nivrofrenate was collected in sparsely 
forested areas and savanna, and purticularly in 
qpen patches of grass on damp or swampy 
eratnd, Same specimens were taken beside air- 
strips and others in grass-covered clearmgs 
bordering tracks through the forests, His an 
extremely timid nocturnal species, leaping 
rapidly towards the cover of dense vegetation 
when disturbed, By day it behaves similarly 
when found sheltering in tall grass. 


Litoria rothi (de Vis) 


Lirorle rorhi is a member of the L. perorit 
species graup. Until recently it was regarded 
as identical with £. perent whieh, bn reality, 
it replaces in north-east Australia (f. R. 
Straughan, personul communication). 


The New Guinea material consists of the 
following specimens: Gnao, Pahoturi River: 
M.C.Z. 84551; Old Zim, Oriomo River; 


M.C.Z, 79727; Wipim: S.A.M. 10623, Boze, 


Hinaliirt Rivers M.C\Z 79728, S$1613-15_ 
PING M. (F.PL) 3350; Weam: M-C.Z. 79729, 
S.A.M. 13113; Morehead: M.C_Z%. (1.P.) 
37605 (3), PNLGM. (F.P.) 561, SAM, 
13111-12. 


This arboreal species is characterised by the 
following features: size moderate (maximum 
stioul to vent length of males = 42 mm, 
females 44 mm); fingers approximately one- 
Murd Webbed with flattened, dilated terminal 
discs;, tympanum partially hidden by  pro- 
nounced supratympanic fold; dorsal surfaces 
of preserved specimens dull brown, sometimes 
with poorly defined, irrepulir patches; back of 
Ihighs black, with two or three uniformly 
shaped, pale cream markings, ‘There ate black 
markings in the inguinal and axillary regions, 
and a black line wlong the inferior margin of 
the supratympanic dermal fold: ivis (in tife) 
dull ted above the pupil, and pale etey or pale 
geld below i. 


The only member of the F, perany spectes 
group formerly known to occur in the southern 
lowlands of New Guinea is 2. anthoinensty, 
reported by Tyler (1968a) from Mabilahol on 
the Ok Sih River in West Friaa, and more 
recently found by one of us (F.P.) to coexist 
with L. rathé in the Western District. Liveria 
amboinensis is a much larger species wilh fully 
wehhed fingers, and lacks the black thigh 
markings of J... reathi. 


Litoria roi was found in sparsely wooded 
areas and in monsoon forest. It was commonly 
collected at night on leaves of plants within 
2m of the ground; the series collected al Rue 
were on the leaves of an ornamentel Cretan 
near village houses. Durmy daylight at was 
tuund within curled leaves, under bark anid in 
similar tefirges- 


Litorin rnbella (Gray) 

Litoria rubella is a wide-ranging species. in 
Australia, oceurring in diverse habitats over 
the entire northern half of the continent. Fts 
presence in New Guinea hus been established 
by the collection of the following 77 speci- 
mens: Daru Island: M.C.Z, (F.P.) 30592: 
Weam: M.C.Z, (VP. 31467, 81609-12, 
§1099-81100, Morehead: A.M.N AL 84527, 
SAM. LOK3O (2), 13720-23, (3125-29, 
P.N.G.M, (F.P.) 334, (F.1) 362, M.CZ 
(F.P.) 30936 (2), (F.P.) 30937 (2), (F.P.) 
30938 (14), (RP) 37606, (FP) 37607 (301, 
ALM. 32709-L4 


HYLID TROG FAUNA OF NEW GUINEA 163 


The characteristic features of the species are: 
small size (maximum S-V¥ of males 32 mm, 
females 34 mm) and squat habitus with very 
short hind limbs; expanded digital discs; fingers 
approximately one-third webbed and toes ex- 
tensively webbed; colouration brown with 
broad, darker brown dorso-lateral stripes on 
the body. In life the dorsal surfaces are brown, 
suffused with a greyish, yellowish or reddish 
tinge. All specimens taken ot Morehewd were 
densely marked with smyll black flecks, but 
such markings were absent in those from other 
localities. The posterior surfaces of the thighs 
are either immaculate yellow or unpigmented 
but for a fine brown stippling. 


Of the species occurring in New Guinea and 


recognised by Tyler (1968a), L. rubella is most 
closely related to L. congertita, from which it 
is readily distinguished by its possession of the 
dark dorso-lateral stripes, and by shorter hind 
limbs (TL/S-V range for rubella — 0.335- 
0.432: 0.477-0,520 for congenita, vide Tyler 
1968a, p. 72). 


Litoria rubella was collected mainly in open, 
man-made grasslands within lightly forested 
country. Most specimens were collected on air- 
strips. It is a nocturnal species, hiding by day 
in situations such as cracks in fence posts and 
amongst plant Icaves. At night it was found 
within 2 m of the ground, and consistently did 
not make any attempt to escape when 
illuminated by a spot light, 


Acknowledgements 


We are greatly indebted to Dr W. G. Inglis, 
Director of Environment and Conservation, 
South Australia, for constructive criticism of 
the manuscript, and to Mr J. 1. Menzies, 


Depariment of Biology, University of Papua 
and New Guinea for specimens of L. timida 
from Brown River Forest Reserve, 


References 


Loverioce, A. (1956)—Rediscovery of Hyla 
dorsalis and Leckrivdus papuanus in New 
Guinea. Breviora ($5), 1-4, 


Macieay, W. (1878).—The batrachians of the 
“Chevert™ Expedition. Proc, Linn. Sac. 
N.S.W. 2 (2), 135-138, 


Moore, J. A, (1961).—The frogs of Eastern New 
South Wales. Bull Amer. Mus, Nat. Hist. 121 
(3), 153-385, 

Parker, F. (1970).—Collecting reptiles and am- 
phibians in New Guinea, Aust. Nat. Hist. 
309-314. 

Rocx, J. (1920) —Note sur la présence du genre 
Crinia, amphibien cystignathide, en Nouvelle- 
Guinée. Rev. Suisse Zool. 28 (5), 115-117, 


TYLer, M. J. (1968a).—Papuan hylid frogs of the 
genus Ayla. Zool. Verh. (96), 1-203, 

Tyrer, M, J. (1968b).—A taxonomic study of 
hylid frags of the Ayla lesneuri complex 
occurring in norih-western Australia. Ree, S. 
Aust, Mus. 15 (4), 711-727, 


Tyter, M, J, (1971).—The phylogenetic signifi- 
cance of vocal sac structure in hylid frogs. 
Univ, Kansas Publ. Mus. Nat. Hist. 19 (4), 
319-360. 


Tytmr, M. J. (in press) —An analysis of the lower 
vertebrate faunal exchange between Australia 
and New Guinea, Jn D. Walker (Bd.), 
“Torres Strait: disjunction or no barrier’. 
(Australian National University: Canberra.) 


VOL. 96, PART 4 


30 NOVEMBER, 1972 


TRANSACTIONS OF THE 


ROYAL SOCIETY 
OF SOUTH AUSTRALIA 


INCORPORATED 


a a 


CONTENTS 


Kott, Patricia The Ascidians of South Australia II. Eastern Sector of The Great 


Australian Bight and Investigator Strait - 


= =a 65 


Dubois, Georges, & Angel, L. Madeline Strigeata (Trematoda) of Australian Birds 
and Mammals from the Helminthological Collection of the Uni- 


versity of Adelaide - - 
OBITUARY: KEITH RODNEY MILES - 
Annual Report of Council, 1971-72 - - 
Award of the Sir Joseph Verco Medal - - 
Balance Sheet - - - - - ~ 


List of Fellows - - 2 5 = 2 


- perl be 


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


THE ASCIDIANS OF SOUTH AUSTRALIA II. EASTERN SECTOR OF THE 
GREAT AUSTRALIAN BIGHT AND INVESTIGATOR STRAIT 


BY PATRICIA KOTT 


Summary 


Seventy-two species of ascidians from South Australian waters are discussed. Fourteen of these are 
new and 42 have previously been reported from South Australian gulf waters. Morphological 
convergence to exploit the environment is evident in many of the forms present. The data support 
the existence of a marine faunal boundary at the eastern end of the Great Australian Bight. It is 
suggested that there are ecological factors responsible for the difference between the South 
Australian gulf and open coast ascidian fauna. 


THE ASCIDIANS OF SOUTH AUSTRALIA II, 
EASTERN SECTOR OF THE GREAT AUSTRALIAN BIGHT AND 
INVESTIGATOR STRAIT 


by PATRICIA Ko1tT* 


Summary 
Seventy-two species of ascidians from South Australian waters afe discussed. Fourteen of these are 
new and 42 have previeusly been reported fram South Australian gulf waters, Morphological conver- 
gence to exploit the environment is evident in many of the forms presént. The data support the exi{s- 
tence of a marine faunal boundary st the eastern end of the Great Australian Bight. It is suggested thut 
there are ecological factors responsible for the difference between the South Avstralian gulf and open 


coast ascidian fauna. 


Introduction 

Collections of uscidiuans have been previously 
made from St. Vincent and Spencer Gulfs 
{Kort 1972a) and from Port Phillip Bay and 
other locations on the Victorian coast (Millar 
1966). The present collections are the first 
reported on from the southern coast of Aus-~ 
tralia between Middleton Beach. Albany (S.W, 
Aust.) and Spencer Gulf. 

Seventy-two species of the Class Ascidiacea 
from South Australian locations (Fig, 60) are 
recorded. Of these. 42 had already been re- 
ported From St, Vincent Gulf, Spencer Gulf 
and Encounter Bay (Kott 19728). Fourteen 
new species are described, 

The association of species in the areas being 
considered differs From that in the gulf areas 
(Kott 19724). Colonies im which each zooid 
maintains independent openings to the exterior 
are upparently favoured and many species 
demonstrate convergence in their adaptations 
to exploit the environment. especially in re- 
yard to their siphonal apparatus. 

The Jarge number of specics taken. and 
especially the large proportion of new species, 
teflects the new habitats that ate now being 
explored by SCUBA diving. 

Type specimens are deposited in the South 
Australian Museum (SAM) or the National 
Museum of Victoria (NMV}, 


Order ENTEROGONA 
Suborder APLOUSOBRANCHIA 
Family CLAVELINIDAE 
Subfamily CLAVELININAR 
Clayelina mirabilis n.sp. 
Type Location: Waldegrave I.: in gravelly 
sand, attached to limestone, 23 m, Shepherd. 


Holotype; SAM, E902, Paratypes; SAM, 
E903. 
FIGS 1, 2. 
Deycription: The colonies consist of a spheri- 


eal, sand covered, base from 2-5 cm in <liam. 


with a thick, naked, branching stalk arising 
therefrom, The thoraces of zooids extend from 
the terminal branches. each ericlosed in its 


own test covering. The living specimen is buff 


or yellow brown, In preservative, however, the 
test of the stalk is reddish-purple, although the 
terminal, free thoracic parts of the zovids are 
almost. transparent. The gravel and sand 
attached to the spherical basal part forms a 
firm outer coat. 

The zooids are up to 4 cm In length and 
extend parallel to one another down through 
the stalk partion inte the base of the colony. 
The wtrisl aperture is sessile although the 
branchial aperture is om a short siphon, When 
the thorax is contracted, the atrial aperture is 
withdrawn to the middle of the dorsal surface, 
while the branchial aperture remuins terminal, 
When the zooid is extended, however, the atrial 
aperture is produced to the anterior end of the 
zooid, more or less Jevel with the branchial 
aperture. There are 15 Jongitudinal thoracic 
muscle bands extending from the ventral sur- 
face, from the branchial aperture and from the 
mid line between the branchial and atrial aper- 
tures. These extend towards the posterior end 
of the thorax, and continue in a wide band on 
either side of the abdomen, There are 18 rows 
of about 45 rectangulur stigmata, The stomach 
is present in the posterior end of the abdomen 
and it has 4 thickenings in its wall, 

Remarks: The zooics of this species are larger 
than other colonial species of the Clavelinidac, 
and the species is unusual in that the zooids 


* Zoology Department, University of Queensland, Qld.. Australia 4067: 


Trans. R. Soc. S. Aust. Vol. 96, Part 4, 30 November 1972. 


166 


extend the whole length of the colony and are 
hot confined to the upper free part as in Pado- 
clavella cylindrica; nor are they completely 
embedded as in Clevelina buucdinensis. (see be- 
low). 

Most species of the family are attached tua 
Substrate by the hottem of the stalk. In the 
present species, however. the unique adhesive 
property Of the sides, as well as the hottorn of 
the basal part of the colony, has resulted in 
un encrustation of sand, shell and olher foreign 
Particles to form its distinetive hard, spherical, 
surface Which appears lo have heen embedded 
in the substrate to anchor the colony. The habit 
iy most unusual since the spherical shape of 
the base of the colony does not appear to pro- 
vide i. rigid attachment or root system and 
there could be some movement of the colony 
in the substrate. 

The habit may be compared with that of 
certuitt Stolidobranchia, und VPhlebobranchia 
where adherent sand forms a hard protective 
envelope around free-living individuals thal 
are not fixed to the substrate. 


Claveling nodula p.sp, 
Type Location: Off Waldegrave 1, Sliep 
herd, Molatype; SAM, E8938, Paratypes: 
SAM, E908. 


FIG, 3 

Description; The colony consists of a fidm, 
translucent. branching stalk of 1.0 cm im diam. 
The zooids, separate [rom one another and 
cach in its independent covering of delicate, 
complelely transparent, lest, are crowded 
around the free ends of the common stalk and 
ts branches. The living zavids are bright 
orange in the centre but colourless in preserva- 
tive, In the present specimens the zooids are 
mostly rctracted into the common stalk, They 
are 4-8 cm long, depending on theit degree of 
contraction. The abdomen is many times 
jomyer than the very short thorax, There are 
up to 20 oblique or longitudinal (sometimes 
coulescing) muscle bands on the thorax, of 
which 5-10 cross the mid-ventral line posterior 
10 the branchial siphon. The thoracic muscula- 
ture extends to the postero-dorsal corner of the 
thorax and continues in wide bands along 
either side of the abdomen, There are three 
rows af ahout 20 stigmata, The gut loop may 
he relatively long. The stomach is in the 
posterior end of the abdomen and its elandular 
liniag Is divided into four sections. 

Remarks; The zooids of this species, although 
smaller. resemble closely those of Pycnecluvella 
diminura (Kow), although there pre fewer 


PATRICIA KOTT 


stigmata in the present species. ‘Ihe colony 
form provides the main distinction between the 
species, In P. dininura zooids are parallel \o 
one another, and muy be entirely independent 
although the test of adjacent zooids may fuse, 
ur be confiuent for varying distunces from the 
base to form a stalk. In Clavelina nodula the 
vouids are not parallel to one another, but 
tadiate from around the tree end of a distinct 
common stalk. Clumps of protruding zoolds at 
diferent points along the stalk precede the 
formation of a branch and appear to contri- 
bute to the later development of that branch. 
The addition of new zooids, developing from 
vascular stolons within the stalk, do not 
directly afleet jis diameter although they are 
accommodated in the stalk when withdrawn 
froin the surface as in the present preserved 
specimens, 
Chlaveliua baudinensis Kott, 1957: 87 (apeci- 
mens with smal) larvae, {rem Rottnest 4. 
and tayerton Bay). ?Millar. 1966: 363- 
Non Clevelina havdinensis Koit, 1972a; 
ae 
New Record, Elliston Bay, 
Records; W, Aust. (Rottnest 
(Laverion Buy)—Kott 1957. 
Dexeviption: Small capitate colony consisting 
of an alinost spherical head and short wide 
stalk. The test of the head is very delicate and 
completely transparent while that of the stalk 
is firmer. Zooids arc embedded completely and 
open jl] around the surface of the head from 
which they radiate in toward the stalk. ‘lhe pre- 
served zooid is transparent with pigment spots 
in the mid line, dorsal and ventral to both 
apertures. There are ahowl 12 longitudinal 
muscle bands in the thorax, of which only 
2 extend across the mid line ventral to the 
branchial siphon, There are 16 rows of ahout 
40 stigmata, ‘The stomach is about half way 
down the abdomen and is small, with its inner 
glandular wall divided into Jour distinct sec- 
tions. 


Previous 
pe) Vie. 


Remarks: A reéexamynition of specimens pre- 
viously identified as this species (Kou 1957) 
hus shown that indeed there are two species 
Tepresented, sugecsted by different larvae 
(Kotte 19690), and that C. haudinensis iz dis- 
unguished by its slightly smaller zooids, with 
invre rows Of sligmaty hut with fewer longitu- 
dina] museles on the thorax und crossing the 
ventral mid line. Zaoids in the present species 
also tend to radiate in from the sutfuce of the 
head in contrast to a more parallel arrange- 
ment observed in othey speciiens that rep 


ASCIDIANS OF SOUTH AUSTRALIA IL 


Tesent fi distinct new species to be described in 
u later Work (Rott, in press (3))- 


Fodoclavella moluceensis Sluiter. Kott, 1972a; 
S and synonymy, 

New Records: Ellision Bay, For Previous 

Records, Description, see Kott 1972a. 
Remarks: The zooids cising from the basement 
metnbrane ore separate from one another for 
thetr whole extent. There are four pigment 
spots dorsal and ventral to and on either side 
of the branchial aperture. 


Podoclavella cylindrica (Quoy & Gaimard). 
Katt, 1927a: 5 and synonyniy. 


New Record; Waldegrave L For Previous 
Records, Descriptions, sce Kott 1972s. 


Podclavella meridionalis Herdman, 1891; 603; 
1899: 4 Hartmeyer. 1919; 14, Hastings, 
19312 81. Kort, 1957: 91. 

New Record: Pearson I, Previons Records: 
W. Aust. (Cape Boileay. Cape Jaubert}— 
Harnmever 1919, Old. (Great Barrier Reef, 
Muckay}—Hastings (931: Kott 1957, 
N.S.W (Port Jackson, Nelson's Buy, Part 
Stephens )—Herdmun 1899. 

FIG. 4 

Deseripion: Only a single specimen ts avail- 

able. This consists of a long stalk and rounded 

head slightly shorter than the length of the 
stalk. The whole zooid is about 10 cm high, 

‘The test ig soft, Wansparent and gelatinous 

on the head but the test of the stalk is tougher. 

The test surrounding the siplions is especially 

delicate, The body wall of the zovid is pig- 

mented blue to black and is.easily seen through 
the transparent test. The branchial aperture is 
terminal and recurved so that the opening & 
directed laterally and downwards. The smaller 
atrial aperture from the antero-dorsal corner 
of the head is directed upwards, There are 
3 rows of branchial tentacles at the base 
ef the branchial siphon, The largest tentacles 
in the mest posterior row allernate with 
inoderute-sized tentacles in the middle row, 
while in the third and most anterior row there 
are twice the number of small tentacles. There 
is no cue dorsal lamina although tianyulur 
lunguets afe expanded from the transverse 
vessels at the side of the mid-dorsal line. There 
are 34 rows of gt least 100 stigmata on each 
side of the body, The dorsal tubercle has an 
undulating longitudinal slit. In the present 
specimen the abdomen is very contracted and 
the course of the gut obsciired. The stomach 


167 


is identified as a region where the gut wall has 
au glandular appearance but does nat appear to 
be of gréater diameter than the rest of the 
intestine. The body wall is thick and heavily 
pigmented. Longitudinal muscles extend from 
both siphons and cross one another on the 
thorax and extend down both siles, of the 
abdomen, 

Remarks: Although records of this species are 
few, tt has a circum-Austtalian discrilhueion_ br 
could be a common component of the fauna 
in sheltered caves and reef overhaiigs where 
hitherto it has been inaccessible to collectors. 


Oxycorynia arenosa m.sp, 
Type Lecation: Invesugator Strait (St, 
¥18). low, flat reef, 30m. J, Wursen, Hale- 
type: NMV, H168. Paratypes: NMV. H169. 


FIGS. 5-7 

Description: The colames form cylindrical or 
finger-like lobes only very slightly expanded at 
their Free end, They are 1.0-1.75 em in diam, 
and up to 9 em long, They sometimes branch 
along their length or rise from 2 common 
base. The surface of the central Jobe is sandy 
to 8 depth of shout 2 mm, although the 
amerior portion of the zooids protrudes through 
the sandy Jayer and is covered hy 1 thin layer 
of sand-free. transparent test. The central pum 
of each Iohe inside the sandy laver of test i 
soft and transparent. In preservative, the zooids 
have black pigment cells in the thorax which 
confer che dark colour seen through the sandy 
lnyer. Zooids open all around the surface of 
the cylindrical lobes for their whole length, 
Zoids extend toward the interior of the colany 
and their long posterior abdominal stolons con- 
tinue down along the length of the lobe and 
into the base of the colony. The branchial and 
atrial openings are plain-rimmed and may be 
funnel-shaped. There are 12 broad longitudinal 
bands of muscle fibres on each side of the 
thorax extending along both sides of the abdo- 
men. There are 6 rows of 2 lang rectangu- 
far stigmaca. The abdomen is about twice the 
length af the thorax with a long oesophagus 
and a spherical stomach in the posterjor end 
of the abdomen The zooid is only about 0.5 
em long. 

Remarks: The small number of rows of stig- 
mata and the arrangement of the zooids radiat- 
ing in Srom all around the eviindrical stalk 
distinguishes this species {rom most others in 
the family where there are usally more rows 
of stigmata aad the zooids open on the free 
ends of Jobes und extend through the colony 


168 


parallel to one another. Vhe parallel tongi- 
tudinal thoracic muscles cxtend from both 
siphons along the length of the body and do 
not, as in the other species of this family, 
extend front across the endostyle. Oxycory née 
fascicularis (Drasche} (see Michnelsen 1930 
for synonymy). does have 200ids radiating 
from around the head of the colony but here 
there is a distinct, zooid-free stalk distinguish- 
meg it from the present species und the zooids 
are completely embedded, 


Subfamily HOLozoinAE 


Atapozaa marshi Brewin, 1956; 31. 
New Record: Investigator Strait (Stns. ¥14, 
17)_ Previens Record; W. Aust. (Trigg 1) 
—Brewin 1950. 


FIGS. 3, 9 

Veveriptions The colonies form lobes of vary- 
ing lengih up to 6 cm and from | to 2? em 
wm diam. The zooid-free basal stuilk 1 more or 
Jess the same diameter as the head of the 
colony, although it does not have adhering sand 
and is of a slightly firmer consistency. There is 
wu very thin encrustation of sand on the surface 
fest around the head of the colony. The zooids 
epen tw the surface by separate, 6-lohed 
branchial and atrial apertures. There is a 
brown pigment spot ubove the dorsal tubercle, 
There are about 15 longitudinal museles on the 
thorax and three rows of 15 stigmata. The 
oesophagus is short. The rounded stomach is 
especially small and is smooth-walled, ‘There 
is a large group of male follicles to the left of 
the gut loop. In one colony there is a single 
immature embryo contained in a brood pouch 
from the postero-dorsal corner of the thorax, 
Remarks: The colony is similar to that of 
Oaveorviria —fayiculuris — (Drysche) (see 
Michaelsen 1930), and the shape of the 
colonies in both species varies in a similar 
lashion. The zooids, however. distinguish the 
specics. 


Atapozoa tivabilis .gp. 
Type Location; Elliston Bay. floor of cave. 6 
m Oher Records: Elliston Bay, 11 m, Shep- 
herd Halatype: SAM, E899. Pararype: 
SAM, E896. 
FIG, 10 
Description: The colony consists of hallow 
lobes and Jumellag cowescing with one another 
to form a thick and convoluted niass about 15 
em long ancl 4 cm high, traversed by passages 
and spaces. There are large common cloacal 
uperiures present, randomly distributed around 


PATRICTA KOTT 


the lobes, und these are often, but not always, 
near the top of the colony. The zonids are 
randomly irrunged ind do not appear tm be 
in rows. They are seen us white dots in the 
semictranspurent anil very soft test. The zooids 
open by their branchial apertures onto the 
outer surface of the colony, while the atrial 
apertures, directed posteriorly, open intn the 
internal connecling cavities of the Iobes aid 
lamellac, Both apertures are 6-lobed and sup- 
ported on siphons of whieh the branchial is 
half the length of the posteriorly directed atrijt 
siphon. There t a short abdomen about the 
same Jength as the thorax and a bulbous 
vasculal’ process extends fram the posterior 
end of the abdomen. This process varies greatly 
in length in different zooids, Zooids are 2-3 
com long. The thorax has a layer of fine circular 
muscle hands continuous arciind the siphois, 
and there are fine longitudinal musele bands 
internal to the circular muscles, These extend 
dewn the thorax and join into bunds alone 
either side of the dorsi! surface af the abdo- 
men and along cither side of the vasculyr 
appendage, ‘There are 3 muws of branchial 
tentacles, and 3 rows of 8 long rec- 
Eingular stigmata, The dorsal lamina is rep- 
resented by 2 pointed languets opposite the 
transverse vessels, There is no sign of the 
gonads in the gut loop in the present specimen, 
The stomach is spherical und amooth-walled 
and is present half-way down the abdomen. 
The oesophagus is fairly long, The rectum is 
often turned over into the proximal purt of the 
posteriorly-directed atrial siphon, which is 
often blown out into what appears to be a 
large bulloon-like reservoir. 
Remiurks: Both the colony and the zooids 
resemble Atapazea deerata (Sluiter) and A. 
vasta (Millar) (sce Kott 1967). The hranchial 
aperlure in the present species, however. is 
not so Jong, the zooids are not protected by 
furrows and ridges of the test a8 i A. deerate, 
amd there js no central mass of test around 
which the cloacnl spaces ramify. since hete 
the centre of cach lobe ix occupied by a large 
common clotcal space, The vascular process 
has also been desertbed previously for species 
of this genus (see Sigillina deerata! Hastings 
(931), 
Ajupovoy fantasiana (Kott), Kott, 19724: 7 und 
synonymy, 

New Recerd: Denial Bay: For Previous 

Records, Description, see Kott 19722, 
Remarks; In one specimen from Denil Ray, 
there is some purple pigment scaltered through- 


ASCIDIANS OF SOUTH AUSTRALIA IL 169 


Figs. 
Fig. 


L, 2. Clavelina mirabilis. Fig. 1—Colony. Fig. 2—Thorax of zooid showing musculature. 
3. Claveling nodula_ Calony, 

Fig, 4. Podoclavella mericdionalis. Individual, 
5 


Figs. 5- 7: Oxycerynia urenosa. Fig. 5—Paortion of colony showing zooids, Fig. 6—Outline whole 
colony. Fig. 7—Zooid. 

Figs. 8, 9. Atapazea marshi. Fig, 8—Colony, Fig. 9.—Zooid showing posterior abdomina] muscu- 
lature, 

Fig. 10. Atapozea mirabilis. Zooid. 

Figs. 11.12. Distaplia distomoides, Fig. 11.—Colony. Fig. 12.—Zooid. 

Figs. 13.14. Pyenoclavella diminuta. Fig. 13—Colony. Fig. 14—Zooid; musculature removed from 


abdomen. 
Fig. 15. Polycitor obeliscum. Zooid. 


170 


out the test and in the other specimen from the 
same station the test Is semi-transparent and 
there are distinct rings of black pigment in the 
surface round each of the apertures. Otherwise 
both are similarly cather fleshy, flat and. invesi- 
ing colonies. and the zooids are identical. 


Distaplia distomoides (Herdman). 
Amaroucium distomoides Herdmin, 1899: 
75, 
New Records: Waldegrave I. Elliston Bay 
Previous Record: N.S.W. (Port Jackson). 


FIGS. 11, 12 

Description: Vhe colonies are cone-shaped and 
supported om a thick fleshy stalk. Living speci- 
mens are tusty-brawn ar brilliant purple, 
ulthough in preservative they are cream to huff 
coloured, The rest is soft and there is no 
adherent sand or fareign particles. Che zoaids 
are atranged in circular to oval systems all 
round the head. The stalk is zooid-free. There 
wre longitudinal to oblique muscles on the 
thorax and there is the usual wide atrial open- 
ing protected hy a well produced anterior sip, 
There are $ longitudinal and oblique muscle 
bands on the thorax. There are 4+ rows of 
wbout 12-14 stigmata crossed by purastignnatic 
vessels, The stomach is shield shaped and 
has 8 rounded ridges internally, The 
oesophagus is relatively short and there is a 
posterior stomach in the descending portion at 
the gut loop. There is a large rosette of male 
follicles fo the right ol Ihe gut loop. As the 
colony becomes larger. the stalk is reduced. 
und the lurgest colonies are almost entirely 
sessile as in Herdman’s type specimen. 
Remarks: The present species  resenibles 
Distuplia yalhi Herdman ¢see Van Nume 
1918) especially in the shape of the coleny, 
and in the reduction in the length of the stalk 
ws the vzooll bearing heal increases in sive. 
It is distinguished by the small number of 
stigmata in each row and Lhe 8 conspicuous 
glandular stomach folds. Herdman’s type 
specimen of This species is redescribed jn Kolt 
(in press (2)), 


Distaplia stylifera =(Kowalevsky). Brewin, 
1953: 60 and synonymy. Kort, 1957: 95, 
Millar, 1963; 713. 

Didemmniun stvlifera Kowalevsky, U874: 
4443, 
New Recard: North of Waldegrave L Pre. 
waus Records; W. Aust. (Cape Jaubert ro 
Fremantle). Qld (Port Termyson)—Ro 


PATRICIA KOTT 


1957, Also the Red Sea, South Africa, ine 

the east coust of north America (see Brewin 

1953). 
Description: The colany consists of a rounded 
head | cm long, on a stalk of Jess diameter but 
approximately equal length. There are 4 
rows of 12 stigmata and the zooids are 
arranged in oval to circular systems opening 
evenly argund the head, Mature gonads are not 
present, 
Remarks; Although the shape of the colony 
and the zooids are tenticul with those of J), 
wylifera, the absence of the diagnostic gonads 
In a Sac separated from the abdomen prevents 
the posilive identification of this single speci- 
men. 


Sycozoa peduncitlita (Quay & Gaimurd), Katt, 
1972b; 234 and synonymy. 
Aplidie pedunculetnin Quy & Gaimard, 
18342 626, 
New Record: Unvestigutor Siruil (Stn. ¥16) 
For Previnus Records, Description, soo Kott 
1972b. 
Remarks; A single small specimen only is 
available. 


Sycozoa cerebriformis (Quoy & Grimard), 
Kott 1972b:8 and synonymy. 
New Records; Denial Bay, near Ceduna. 
Elliston Buy. For Previony Recordy, Desevipe 
tian, see Kou 19728. 


Family POLYCITORIDAE 
Pvenockivella diminuia (Kotl), Millar, 

715. 

Clavelina dimimuda Kou, 1957: 89, 
New Records: Elliston Bay. Spencer Gulf 
(Tipura Reet), St. Vincent Gulf (afl Por 
Noarlunga, 15 m depth, on rock or epizcic 
On other aseidiuns). Previeus Records: W, 
Aust. (Cape Boileau, Rottnest I[.)—Kott 
1957: Millar 19863. 

FIGS, 13, 14 
Deseriptian; Colonies are 2 em high. Two or 
more zooids muy be fused basally but 
anteriorly the thoraces.of the zooids are always 
independent. The test is semi-transparent 
throughout and contains spherical, dark bodics, 
especially anteriorly, There are 3 rows of 
stigmata and about 12 fine longitudinal muscle 
hands on the very short thorax, extending along 
the ventral surface of the abdomen. ‘The abda- 
men ts about twice the length of the thorax. 


1963: 


ASCIDIANS OF SOUTH AUSTRALIA II 71 


The stomach, in the posterior end of the abdo- 
men, is sounded and smooth-walled externally 
but there are some longitudinal interruptions in 
the glandular wall. Some of the thoracic muscle 
bunds extend across the endostyle while the 
more dorsal bands extend from the atrial aper- 
ture and from across the mid-line between the 
apertures, The specimens are identica) with 
the [ype specimens from Rottnest L and are 
larger than those from Cape Boileau (Millar 
1963}. 

Further colonies, apparently of this species, 
were collecied from Vipara. Reef in Spencer 
Gulf (Shepherd. th m., 20.viii 1971). The 
living zooids are yellow, As in Millar's speci- 
mens, the zooids ure only | cm high, of which 
the upper one quarter is clear glassy test with 
the usual enclosed dark spherical bodies. The 
remainder of each zovid is encrusted with 
sand and is adherent to adjacent zooids, 
Basally, the test tapers into a fine roet-like 
stolon with fine side branches and the basal 
part of the colony is 4 tangled mass of these 
stolons. There docs nol appear lo he any 
organic conlinuily belween the stolons of adja- 
cent vooids and there is no basemen! mem- 
brane as in previously described specimens. 
The zooids have only S§ thoracic muscle 
bands which exlend along both sides of the 
abdomen in fine bands; and aljhough there are 
the usual 2 rows of stigmata, there are 
only 16 sthomala in cach half row, Num- 
bers of beth muscle bands and stigmata are 
therefore much reduced in these specimens 
from Tiparu Reef, The stomach is of the usual 
form, Embryos start their development at the 
base of the @viduct as is characteristic of this 
genus. Well doveloped embryos taken from 
the oviduct ahout half way up the abdoinen 
are 1.3 mm long, the tail is wound once around 
the body, there is un ocellus but no otolith, and 
there une three “tube-like pupillae charac- 
teristic of the genus (Trason 1963). 


Remarks. All specimens share the pycnoglave- 
hd characters of short thorax and Jarge eggs 
which are fertilized at the base of the oviduct, 
developing as they pass up toward the atrial 
apertire, The absence of the basal membrane 
in the specimens from Tipara Reef could be a 
response tn the sanidy substrate in which they 
are rooted. There is considerable variation, 
however. in the number of stigmutu and the 
number of longitudinal muscle bands in the 
specimens, which ix not related to the size of 
the zoaid, Further collecting may demonstrate 
that more than a single species is involved. 


Polycitor giganteum (Herdman). Kolt, 1972a: 
9 and synonymy 

New Records: Waldegrave L., Elliston Bay, 

Pearsun J. Investigator Strait (Stns, X15. 

21). Previous Records: See Kon 1972x. 
Beseription: The present colonies vary {rom 
small, conical and sessile, to large and spheri- 
cal, constricted from a sandy base. The basal 
test is translucent but the test of the head is 
almost glassy and transparent. Zoolds open all 
round the head and radiate inta the buse of the 
colony as is usual for the species. There are 
15 Jongitudinal thoracie muscle bands extend- 
ing in w wide band along the ventral half of 
the ybdomen, There are 10 rows of abour 20 
stigmata. The stonvach, in the posterior end of 
the abdomen, has four folds. Gonads ure 
present in the gut loap. 


Palycitor obeliscnm n.sp. 
Type Locality: Investigator Strait (Stn, 
YIS), 30 mi oon aw low, fat reef, Marsan, 
Holotype: NMYV, Ht67, 
FIG. 15 
Description: The colony forms a pointed, 
sessile cong, The test is gelatinous and firm, 
und there is sand basally. Zooids epen all 
around the surfuce and appear to he arraneed 
more or less in longitudinal bnes. Zooiws 


_tadiate in from the surface to the base of the 


colony. The abdomen is about four times the 
length of the thorax, Both apertures ure 6- 
lobed and the atrial aperture is on a short 
siphon, There are 20 longitudinal thoracic 
muscles and some transverse muscles on the 
thorax, There are 5 rows of about 12 stig- 
miata. The stomach which js present in the 
posterior third of the abdomen is large and 
smooth-walled although it may be collapscel 
into folds. There is a small, rounded, posterior 
stomach, 

Reriarcs: A colony of this species superficially 
resembles that of Distaplia distomoides, The 
zooids, however, are typically of the genus 
Polyciror and are distinguished from other 
species in that genus by the very small number 
of raws of stigmata. 


Eudistama, reniert (Hurimeyer). Koll, 1972a: 
[TQ and synonymy, 
New Record: Elliston 
Records: see Kot |972a, 
Description: Vhe present colony forms an 
nregularly elongate wr oval to circular cushion 
with rounded walls. fe is op to 1,5 em high 


Bay,  Previnns 


172 


and 3 cm in diam, itis fixed by most of the 
busal sirface and the upper surface is smooth. 
In the living colony, the circles of zowjids show 
as bright ted stars in a pale test, However, 1) 
preservative. the stars are colourless and the 
test is blavk, the pigment being contained in 
the round cells in the test. The zooids are 
arranged in circles of up Lo 6, with the atrial 
openings adjacent to one another in the centre 
of the circle forming a pseuclo-cloacal open- 
ing. There are strong longitudinal muscles on 
the siphon but there is no conspicuous circular 
sphincter, There are 13 strong mmisele bands 
on the thorax extending along either side of 
the ahdomen. There are 15 stigmata in each 
of the three rows. 

Remarks, The arrangement of the zoolds in 
circles. is usual for this and other related specics 
of the genus, The fleshy firm consistency of the 
colony is typical of the species. 


Cystodyles dellechiajei (Della Valle). Kott, 
1972a: 11 and synonyeny, 
New Record: Elliston Bay. For Previous 
Records, Descriptian, see Kott |972a, 


Family POLYCLINIDAR 


Subfamily FUNEROMANTINAE 
Euherdmania australis Kott, 1957: 103, 
New Records; Elliston Bay, off Waldegrave 
1, Trvestigator Strait (Stn. Y19). Previous 
Records; Vic. (Port Phillip Heads). N.S.W, 
(Camden Haven)—Kott 1957. 


FIGS 16-18 
Description; The colomes are formed of the 
usual sandy, finger-like. Iobes containing a 
single zocid. The atrial aperture ix sessile in 
the middle of the obliquely flattened to con- 
cave free end of the Iobe. The branchial aper- 
lute is just ventral to the flattended tree end 
and is protected above by it erescentic fap of 
sand-stiffened test which covers the opening. 
The aperture itself is in a sand-Free area 
covered hy this flup. There are 13 rows of 
stigmata with parastigmoatic vessels. Branchisl 
papillae ure present in the middle of the pri- 
mary and parastigmiatic transverse vessels on 
each side of the body. The anus is present half 
way up the thorax and has 10 pointed lobes 
on the border. The stomach is stnall, with 
about 16 rather irregular and often branching 
folds, In the specimens from Waldegrave 7. 
ubout § developing embryos are present in 
the Jhorecic purt of the ovidiict, the most 


PATRICIA KOTT 


mature embryo being present toward the distal 
end of the duct. The eggs are therefore fer- 
tilised at the base of the oviduct and appear to 
start their development as they move up toward 
the opening, Testes are bunched in the short 
posterior abdomen, Muscles are present, 
especially round the dorsal border of the 
branchial aperture, but do riot extend far 
down the thorax, When the anterior part of 
the zootd is contracted, the Jower part af the 
thorax is pulled upwards, placing the opening 
of the oviduct opposite the atrial opening. 
Larvae: The larvae are 0.9 mm long. They 
have 2 shallow, wide, papillae in the mid-line. 
Sets of 3 median ampullue are present in the 
intervals between the papillae, and in cach set 
the middle ampulla is farger and its free en 
is Aattened, while the dorsal and ventral am- 
pullae in each set are smaller and conical. 
Small vesicular cells are supported sll over the 
body wull of the larvae, 

Remarks: The larva is of the polyclinid type, 
developing ampullary vesicles ancl with the 
usual papillary cells surrounded by acecssory 
cup-like suckers. The papillae are not modified 
as in other species of this and the related 
genus, Plarentela (see Kott 19694), 


Ritlerella herdmonia Kott. 1957; 102, 1972a; 
11 and synonynvy. 

New Recerd: Elliston 

Records: sco Kott 19724, 
Descriprion; The colomes consist of small 
slender lobes. 1 cm long, with expanded spatu- 
late tips, joined basally, Minute zooids open 
wound the border of the lobes. There are 5 
rows of sligmala and 5 stomach folds Papillac 
are present on the transverse vessel. 


Bay, Previous 


Pseudodistoma australis Kott, 1957! TOL) 1963: 
7S, 
New Records: Waterloo Bay. Waldegruve 1. 
Previnus Recerds: W. Aust. (Rottnest [,)— 
Kott 1957, 1963. 


PIG, 19 
Description: Colouwies are soft, rounded and 
scasilc, up to 2 cm in diam, and no more than 
0.5 cm thick. The test ms especrally soft and 
semi-transparent, The zooids may be orange- 
brown with flecks olf black, and in the pre- 
served colony spherical black pigmient-celfs 
remain, Zooids open over the upper surface by 
two separate, 6-lobed openings. There are 3 
rows of about 2 stigmata, There fre 14 
strong longitudinal muscle bands on the thorax 


ASCIDIANS OF SOUTH AUSTRALIA il 


which extend as a single band down the ven- 
tral side of the abdomen and onto the posterior 
uhdomen, The stomach is small and smooth 
walled. with a distinct typblosolar line. and is 
present half way down the abdomen. The 
oesophagus is fairly short. The thoracic 
muscles extend down the ventral side of the 
sbdomen and to the left of the intestinal loop 
onto the posterier abdomen, which, in these 
specimens, appears to arise fram the left side 
of the intestinal loop owing to the strong con- 
traction of the body musculature. The thorax, 
abdomen and posterior abdomen are of equal 
length. Gonads are not developed in the pre- 
sent specimens, In specimens from Waldegrave 
1, there iy occasionally a large halloon-like 
hrood pouch, 

Remarks: The specimens have been compared 
with the type specimen of Pseudedistoma 
australis and found to be identical, although in 
the 1ype the posterior abdomen ts better de- 
veloped with mature gonads. Psevdodistoma 
cyrnusense Pérés, 6952, and P. fravills 
Tokioka, 1958, are related species ‘with a 
sessile colony, short abdomen and oesophagus, 
amall smacth stemach, and the testis follicles 
bunched at the pastevior end of the pasterior 
ahdomen, Po cyvrnusense has only 12 stigmata 
pet row, while both P. australiv and P. jrapilis 
have 20 or more stigmata per raw. P, fragilis 
is distinguished from the present species by the 
presence of up to 3 embryos in a broad paich, 
While in P. castraliy only a single embryo has 
been found in the brood pouch that is con- 
stricted from the posiero~lorsal aspect. of the 
thorax (see Kott 1963). 


Pseudodistoma cyrnusense Pércs, (952: 37. 


New Record: Elliston Bay. Previous 
Reeords; Mediterranean—Peérts 1952, 
FIG, 20 


Description: The colony is a very soft. irreyu- 
larly rounded, inverted saucer shape, & cm in 
diam, and about I em high In the centre. The 
surface is very smooth and there are no ads 
herent or included foreign bodies. The test is 
shahtly transparent, cloudy and of creamish 
colour. Zooids open separately to the exterior 
over the upper surface, The siphons are fairly 
muscwar and there are 12 fine Jongitudinal 
muscle hands on the thorax gxtending along 
both sides of the abdomen and posterior abdo- 
men, There are 3 rows of stigmata with about 
{2 stigmata pec row, The abdomen is slightly 
longer than the thorax, although the pasterior 
abdomen is about 2 times the length of the 


173 


abdomen, ‘The stomach is half way down the 
abdomen and externally is smooth, althoygh 
internally there appear to be 4 glandular ndges, 
There are minute mulberry-like cells tn the 
common test, from 0.01—0.02 mm in diam. 


Remurks: Tt has not been possible to separate 
the present colony from Pérés’ Mediterranean 
species, The colony form and consistency, the 
numher of stigmata, and the size and shape of 
both the stomach and of the zooid are iden- 
ticul. Pseudodistama aurea ( Brewin 1957) and 
P. manritiana Vasseur, (967, both form large 
fieshy colonies, However, in these forms, the 
abdomen is very much Jonger than the thorax 
und the oesophagus is especially long. Psendo- 
distoma fragilis (Tokoika, 1958) and P, aus- 
irglis also form fleshy investing colonics. but 
they are much thinner than the present speci- 
men and have 25 stigmata in cach row, distin- 
guishing them from the present speciimen 
which has only 12, The present colony is 
considerably more extensive Lhan specimens of 
P. australis so far available. 


Pseudodistoma cereymi Michaelsen, 
19724; 12 and synonymy, 

New Keeords: Ejliston Bay, Waldegrave I., 

St, Francis 1. For Previous Records. Des- 

cription, sec Kott 1972a. 

FIGS, 21, 22 

Remarks: The species is distinguished from 
other stalked forms by the relatively large 
number of theracic muscles (30-40). The 
stomach is rather capacious and has ils internal 
glandular wall divided into four sections (see 
Michaelsen 1924), This also helps to distin- 
guish the spectes [rom P. australis i which the 
stomach is especially small, shallow and smooth 
walled 

The present colonies from Waldegrave J. are 
long, with a Jong stalk and head, while those 
from Elliston Bay are no more than 2 ¢m 
high with a round soft head on a short thick 
stalk. This Tange in the torm of the colonies 
has been Observed in specimens fram New 


Kott. 


Zealund (see Sigillinaria novae-celandiae 
Brewin, 1950, a synonym of the present 


species, and P. cereum; Brewin 1958). 


Placentela ellistani n.sp. 
Type Lovglitv; Elltsion Bay, inside caves, 
outside bar, 14.v-1971, Shepherd. Holorype: 
SAM, E901. Pararypes: SAM, B90) 
FIGS. 23, 24 
Description: The colonies form narrow, fin- 
shaped Johes, rounded at the free cdge and 
narrowing 10 the base, The lobes wre 5 em will 


174 


PATRICIA KOTT 


. Euherdmania australis, Fig. 16—Individual. Fig. 17. —TIndividual removed from test. Fig. 


18—Larva, 
Pseudodisiama australis. Zooid. 
Pseudadistoma eyrnisense. Spicules from test. 


. Pseudadisioma cereum, Fig. 21—Colony, Fig. 22.—ooid. 
. Placentela ellistoni, Pig, 23.—Calony. Fie. 24.—Zooid. ; 
. Polyclinuem nepteninm, Fig. 25.—Individual. Fig. 26—Portion of branchial sac showing 


papillae produced from transverse vessel. 


- Apliditn colelloudes. Fig. 27—Colony, Fig, 28—Larva, 


Aplidiium pantherinum, Thorax. 
Aplidium flavlineattr. \atva, 
4Aplidinm elatum, Zooid. 
Synoieium papilliferium. Zooid. 


ASCIDIANS OF SOUTH AUSTRALIA U 


and aboul 2.5 em broad across the free end. 
which jx the widest par. of the colony. There 
are sand-covered swellings projecting back 
from the rounded Free border of the fun, and 
overlapping both sides of the colony. The test 
is Very stf and sandy. Branchial apertures are 
arranged in an are protected by the overlap 
from the rounded Free border of the fan, Each 
aperture is sessile and in a sand-free area of 
lesl, The atrial apertuircs on the opposite side 
of the colony are in a groove slightly further 
back than the corresponding are of branchial 
upertures. Each atrial aperture is on a small 
mound and, us with the branchial apertures, is 
covered by the overlapping rim of the rounded 
free border of the fan. Zooids are arranged in 
only a single laver at the top of the colony but 
as they extend down towards the base, they 
overlap und cross One another and here the 
colony is narrower but thicker to accommodate 
the posterior ends of the zooids. The 4 lobes 
around both apertures are minute and very 
pointed. There is a mesh-work of circular 
and. longitiidinal muscles on the thorax, extend- 
ing up around the siphons. There is a wide 
musele band on the left side of the abdomen 
and on the dorsal side of the posterior abdo- 
men, formed by very fine lonaitudinal bands 
from the thorax which all cross to the left side 
of the body across the postero-ventral part of 
the thorax. There are 17 rows of about 35 
reclangular stigmata. The oesophagus is very 
short and the stomach is smooth and shield- 
shaped, The ubdomen is only about half the 
length of ihe thorax. The posterior abdomen 
ts Of similar length and contains a large 
number of male follicles bunched together 
posterior to the ovary. It appears from the 
specimens cxamined that the branchial aper- 
jure may be contracted back along the ventral 
surface of the thorax so that the atrial siphon is 
derminal, Thus. by virture of the strong body 
musculature, the whole top rim of the colony 
probubly moves up and down or from side to 
skie according to wheiher the arc of branchial 
apertures or the arc of atrial apertures is to be 
opened af whether both are simultaneously 
to be opened or closed. 


Remarks: The specics is unusual and although 
the arrangement of zooids within the colony 
resembles that found in Ritrerelia hierdmiania, 
the zooids themselves are distinguished hy their 
strong thoracic musculature, by the amooth 
stomach (which resembles the type found in 
the genus Syreiciwin), and by the largce num- 
bet of rows of stigmata. 


Subfamily POLYCLINING& 
Polyclinum neptuniuo: Hartmeyer, 1912: 331. 
Rott, 1963: 83 and synonymy, 

New Records: Elliston Reef. Elliston Bay. 

Previnus Records: W. Aust (Shark Bay ta 

Atbanyj—Michaeken 1920; Kore 1963, 8. 

Aust. (Reevesby [.)—Kott 1963. South 

Africa—Hartmeyer 1912, Millar 1962, 

FIGS. 25, 26 

Deseripiick; Colonies of large investing sheets 
or sessile, almost hemispherical, lobes, + mm 
high in thickest part and from 2 cm in diam, 
There is a dense sandy coat externally, absent 
only where branchig! ppertures open to the ex- 
terior, The common cloacal apertures are 
present on the surfyce. on or at the side of 
sligat rounded elevations, Zooids are in round, 
to elongate or long, double row systems along 
either side of shallow, narrow, common 
cloacal canals. There is no sand in the internal 
test, Which is semi-transparent and <ofe but 
tough, There are sometimes. but not always, 
black spherical pigment cells in the body wall. 
especially in the atrial languet and around the 
branchial aperture. There are 6 small, very 
pointed branchial lobes, The atrial tnnguet, 
rising from above the sphincter muscle. may 
be pointed or Jong and with a flat terminal 
border fringed with up to 7 minute, pointed, 
lobes, The shape of the atrial langue is 
associated with the location of the zooid in 
relation to the cloncal canals or with the 
comman cloacal openings, and its tip may be 
incorporated in the border of the opening. 
There may be 4 minute papilla From the body 
wall below the aperture. There are 6 paral- 
Jel. Jongitudinal muscle bands in the atrial 
fanguet alihough these may coalesce or divide 
at uny point along their length. These are 
crossed by Very fine traverse muscles, There 
are 10 Jongitudinal muscles radiating from 
the siphons and extending down the thorax, 
although they may be difficult to detect in the 
postenor half of the thorax. There are 10) rows 
of (6 stigmata and a similar number of flat, 
rounded lobes, confluent at their base, sup- 
ported along the transverse vessels. “The 
stomach ts small and smooth externally hul 
with glandular papillae internally. The pos- 
terior abdomen ts long and tongue shaped. 
Remarks: The relationships of species within 
this genus have always been difficult ta ceter 
mine, oWing to the homogeneity of the zooids 
and variability of the colony form and atrial 
languets, The present specimens have heen 
identified by the relatively Jarge oumber of 


Ff 


branchial papillae, by which the species ts 
distinguished from FP. macrephyllum (see 
Michaelsen 1930). There ate also fewer rows 
of stigmata than in P. macrophyllum, although 
the number of stigmata in each tow is greater. 
The internal test is also tougher than is usual 
for this genus. 


Aplidium Jobatum Savigny, 1816: 182. Katt, 
1963: 97 and synonymy. Tokioku, 1967: 
22. 

New Record: Elliston Bay. Previour 
Records: sce Kott 1963; Van Name 1954. 
Deseription: Specimens are almost spherical 
and | em in diam, The lower half is sandy and 
there is a more or less flattened upper surface 
through. which the orange zooids are clearly 
seen through the transparent test, Spurse sand 
is present throughout the remainder of the 
fest, The z00ids ure very small, with a fleshy 
tripartite. atrial Janguet from the upper border 
of the opening, There are 6 rows of about 
10 stigmata and 5 pronounced stomach folds. 
Remarks: The small zooids, the atrial aperture, 
the small aumber of stigmata and 5 stamach 

folds characterise the species. 


Aplidium colelloides (Herdman). Kott (972a; 
15 and synonymy. 


New Records: off Waldeerave L. Investi- 
gator Strait (Stn. ¥21), Previous Records: 
sec Kott 1972a. 


FIGS, 27, 28 

Description! There is a rounded, firm, gelat- 
tnous head supported by a tough leathery stalk 
about 17 em Jong. Zooids are minute and 
arranged akong both sides of narrow, branch- 
ing, longitudias] canals, Large common 
cloacal openings are randomly distributed over 
the head. The languct from the anterior border 
of the atrial aperture is very small and pointed. 
There are 16-18 rows of 10 elliptical stig- 
mati ant 135 stomach folds, In the present 
specimens, there are 2 embryos contained in 
the posterior part of the peribranchial cavity. 
One is almost mature while the other is at an 
early stage of development, This difference in 
the stage of development of the embryos is 
observed in all the zooids tn which embryos 
ate present, 

Larvae; Mature larvae are large, L.5 mm Jong. 
There are small, crowded, epidermal vesicles 
projecting from the anterior part of the Jarva 
around the base of the papillac. and extending 
along the yentfal surface. 


PATRICIA KOTT 


Aplidium pantherinam (Sluiter). Kott, 1963: 98 
and synonymy, 


Poammaplidium  pantheritum  Sluiter. 
1898; 26. 
New Record: Elliston Bay. Previous 


Retords: W. Aust. (Rottnest 1. ta Hamelin 
Buyj—Kott 1963. 8. Africa—Sluiter 1893; 
Millar 1955, 1962. 


FIG, 29 


Deseription; The colony is oval in outline, 2 
em thick and 3 cm long. Jt 4s fixed by i small 
part of its base. The base and walls are even 
and sandy. The surface is marked off into 
irregularly circular depressed areas that are 
often free of sund, These are about 5 mm in 
diam, and are separated from onc another by 
raised sandy ridges about 0.5 em wide which 
form a network over the surface and sharply 
overhang the periphery of the depressed areas. 
Common cloacal apertures are present in the 
centre of these depressed areas, and wre sur- 
rounded by the branchial openings of the 
zooids. There 18 sand enclosed throughout the 
otherwise gelatinous test, thus creating a rather 
hard colony. Zovidy »re minute and crowded 
and radiate down into the base of the colony, 
The branchial aperture is surrounded by 6 
well-defined lobes: The jatrial aperture is on a 
muscular siphon fram about half way down 
the thorax, protected by a long, pointed, lip 
tising from the body wall anterior to the 
siphon. The thorax is nmuscular with a well 
developed circular sphincter muscle at the base 
of the branchial aperture. ‘There are. ahout 20 
very fine longitudinal muscle bands on the 
thorax. There are 16 rows of 6 stigmata. The 
abdomen is about the same lenath os the 
thorax, with the small atomach half way down 
the ubdomen, The stomach wall has 5 distinct 
folds. 

Remarks: The form of both colony and 
zooids is unusual, but docs resemble Apliaiunr 
cratiferunt Sluiter, 1909; Van Name, 1918, 
from the Philippines, which is distinguished 
however by its 10-12 stamach folds, 


Aplidiom rubricollum Kott, 1963: 103; 1972: 
15, 


New Record: Pearson J, For Previous 
Records, Description, see Katt 1972a, 


Aplidium flavolineatum (Sluiter). Kott, 1963: 
165 yrd synonymy, 


Amuarouciam fiayolinedtam Sluiter, 1898: 
MM), 


ASCIDIANS OF SOUTH AUSTRALIA IJ 


New Records; Ellision Bay, off Waldegrave 
1, Previous Records: W. Aust. S. Aust, 
Vic,, NS W.—Kott 1963. S. Africa—Sluiter 
1893; Michselsen 1934; Millar 1955; 
Tokioka 1959, 


FIG. 30 

Description: The colonies form low, rounded, 
cushions about | cm high and 7 em in diam.. 
with a sandy basal half sometimes narrowed 
to a short thick stalk. In the preserved speci- 
mens the yooids are orange, and open onto the 
transparent upper surface. They ate afranged 
in circular to oval systems of 5-12 zooids. 
There may be some sand in the surface test 
between the systems. There are 3 pointed 
languets from the upper border of the atrial 
opening. There are about 20 Jongitudinal 
muscles on the thorax, continuous along bath 
sides of the uhdomen and posterior abdamen. 
There are 10-15 tows of 12-15 stigmatu. The 
stomach ts small and rounded with about 25— 
30) very narrow folds. slightly oblique and 
extending antertorly teward the mid Jinc on 
both Jateral and mesial aspects of the stomach, 
The ahdomen and posteriot abdomen are of 
equal length, although the thorax is smaller, 
Zooids. are very small and usually so not 
exceed 4 mim. 

Larvac; These are present in the peribranchial 
eavily, They are 0.8 mm long and the tail 
winds three-quarters of the way around the 
body, There are median ampullae between the 
3 suckers with lateral branches from the base 
of cach median ampulla. The medium ampullae 
ure not always paired as they have been des- 
cribed previously, nor are there posterior 
vesicles. There has been some variation ab- 
served in the form of thts larva however (see 
Kott 1963} and the differences are not thought 
to be significant. 


Aplidinn elatum. a.sp. 
Type Locality: Elliston Bay, outside bar, 
very strony surge, 17 m, 12.1,1971, Shep- 
fwrd. Ficioivpe; SAM E906, Purarypesi 
E905, 
FIG. 31 
Destriprion, The colonies fori tall, unidulat- 
ig, faneshaped, flattened lamellae and lobes 
which are sometimes fused. Each lamella is a 
inuxtimum of 1 cm thick, The maxingum height 
From the buse to the free rounded border is 
6 cm, The surface is sandy but marked off into 
slightly prominent. rounded, swellings corres 
ponding to the anterior ends of ~he minute 
zooids, which open on doth surfaces and on 


177 


the free edge of the lamellae. The test is 
sundy throughout and the colony firm and 
hard, Common cloacal apettures ate present 
from place to place over the surface and 
zooids are arranged in double tows radiating 
from them. The zooids and especially the an- 
tetior part of the thorax and the endostyle are 
crange in the preserved specimens. Zooids are 
about 3 mm long. ‘The thorax and posterior 
abdomen are of about equal length and slightly 
lonver than the abdomen. The branchial aper- 
ture is terminal with 6 sharply pointed lobes. 
The atrial aperture is opposite the 4th—-6th 
row of stigmata and its anterior lip is produced 
into 3 almost foliated lips. These are not 
always of equal size. but they are always very 
muscular with longitudinal bands extending 
along their Jength. There ate about 12 fine 
Jongitudinal thoracic muscles. There are 8 
rows of 8 stigmata. The stomach is very short 
with about 15 distinct folds, 

Remarks: The colonies are very like A plidium 
solidum {Herdman} (see A- arboraramn Kott 
nom, nov, 1963), The species differ however 
in the number of stomach folds, in the length 
of the posterior abdomen, and in the length 
and form of the atrial Jangucts. The colony 
and the atrial lobes are stmilat to those of A. 
sarasinoruny Millar, 1962, from §. Africa. 
However, the body musculature and stomach 
folds differ. Aplidium muliplicarum (see Kote 
(963) forms jelly-fike to firm, investing, 
colomes and has minute zooids, a hranchial 
sac Teserobling that of the present species and 
ubout the same number of stomach folds. 
Again, however, the mitisculur large atrinl 
languei and dense sand inclusion, distinguish 
the present species. 


Synoictum = papilliferum {Michaelseni_ Kote, 
1972a: 16 and synonymy, 
New Record: Waldegrave [, Previoss 


Records. see Katt 19722. 

FIG, 32 
Desermion: The colony is rounded, Jobed and 
branched, The diameter of a single lobe is 
about 1.0 cm. The zooids are parallel to one 
another and at right angles to the surface all 
around the colony. which does. not appear to 
be fixed. There is a sparse coating of sand 
grains on the surface and throughout the test. 
The atrial aperture is on a short muscular 
siphan and protected by a long pointed lip 
which is produced from the anterior border of 
the opening. There is the «sun}) protuherant 
papilla from the body wull posterior cto the 
atrial siphon. ‘There ate 8 fine longitudinal 


178 


muscles 01) the thorax and there are 9 rows af 
VW stigmata, The stomach ts the usual shietd- 
shape and has mulberry-like glandular swell- 
ings. 

Remarks: The form of the atrial siphon. the 
Pupilla and the stomach with tts mulberry-like 
swellings ure characteristic of the species. 


Fumily DIDEMNIDAE 
Tritidemanm cerebriforme Hartmeyer, 1913; 
149. Kott, 1962: 275 and synonymy. 


New Records; Elliston Bay, Investigiter 
Strait (Stn. X17). Previous Recerds: see 
Katt (962, 


FIG. 33 

Desevipilon, There are very extensive posterior 
abdominal common cloneal caviticy and 
secondary conunon cloacal canals at the 
thuracic level. There is a very Lhin basal layer 
of test. Zooids are suspended between the 
busal and surface layer of test by pillarlike 
strands in which the abdomina are embedded, 
and through which the thoracic secondary 
cloacal canals extend, Spiculcs are sometimes 
evenly distribuled throughoul the test although 
they may be thick im the surface layer but 
sparse below thoracic level. They are large, 
0,03-0,06 mm in diam. with 5-7 conical 
potted ruys in optteal fransverse section. 
Zooids have a. minute thorax, with 3 rows of 
sugmata amd a wide atrial opening. There are 
73 coils of the vas deferens around a single 
tests Follicle. 

Remurks: The species is identified by the ex- 
tensive posterior abdomingl cloacal sysiem and 
by the open atrial aperture rather than a pos- 
teriorly directed siphon, usually associited wath 
this type of cloacal system. 


Trididemnum spiculatum Kott, 1972a; 16. 


New Record: Elliston Bay. Previous 
Recards: sce Kotr 197 2a, 
FIG. 34 


Description: The colonies are white and invest 
stalks anc leaves of seaweeds. Deep primary 
cloacal canals extend around clumps of zooids 
bul the secondary canils remain at thoraci¢ 
tevel, There is a long lateral organ, There are 
3 rows of stigmata and 54 coils of the vas 
deferens around a single testis follicle, The 
spicules are stellate, with 5-7 rays in optical 
section, 0.01-0.03 mm in diam. 

Rentarks: The smaller stellate spicules and the 
absence of a large posterior abdominal cloucul 
cavity disthnguish the species. 


PATRICIA KOTT 


Polysyneraton magnilurvam Millar, 1962: 163. 
New Record: livestigator Strait 4Stns MVS, 
YO). Previowy Records; Wytal—Millar, 1962. 


FIG, 35 


Deseription: The colonies are irregularly lobed, 
lurge and Neshy, and are supported by a very 
short and relatively marrow stalk, Each lobe 
may be up to 2.¢m in diam, and a maximum 
uf 2.5 ¢m in length. There are no spicules. 
There i a surface layer of bladder cells and 
beneath this some pigment cells which become 
less frequent internally although they congre- 
gale around inclusions und parasites in the 
test. The zooids are confined to a thin layer of 
surface lest about | mm thick, and the centre 
of cach lobe consists of gelatinous. firm, test 
withoul zooids, The consistency of the colonies 
varies from firm unt gelatinous to hard and 
Lough, but if is thouvht that this may reflect 
the preservation ef the specimens. Common 
cloacal apertures ure rqandomly placed over the 
surface and zooids are arranged on cither side 
of long cloacal cunals radiating from ithe 
apertures. The surface of the colony is marked 
by these long brinching canals in the surface 
layer of test. The zooids are minute und have 
w long oesophageal neck, The thorax is 
especially small, O.S mm Jong. The branchial 
aperture has the usual 6 pointed Jobes and 
there is a wide, open, atrial aperture. There are 
4 rows of @sugmila. There ore § testis follicles 
with 24-54 coils of the vas deferens. The 
ventral surface of each zovid is embedded in 
the common test so that the surface of the 
preserved colony is marked into small rownded 
mounds surrounded by a narrow depression 
Where the thin surface test is depressed over 
the common cloacal canal to which the atrial 
apertures are exposed. 


Remarks: The species is distinguished by its 
Nleshy colony und hy the large number of 
testis follicles and small number of vas 
ueferens coils. Palysyncraton — axpiculatum 
Tokvida, 1949, forms flat investing colonies 
and is often without spicules, but has a long 
bifid atrial lip which is absent in the present 
species. 


Polysyneraton paradoxunt Notl, 1892; 318, 


New Record: Elliston Bay. 
Record: New ZFealand—Nott 1892. 


Previous 


Descripiion: The living colonies are brilliant 
orunge hut in the preserved specimens only 
streaks of orange remain on the surface. There 


ASCIDIANS OF SOUTH AUSTRALIA Il 


are stellate pigment cells scattered amongst the 
spleules. There is a surface layer of dladder 
cells, then a dense layer of spicules which be- 
come less dense in the oesophageal region of 
the zooids and arc absent completely (rom the 
test at the abdominal level and in the basal 
lesl, The common cloweal canals ate very 
shallow and thoracic. There are 8 stigmata per 
row, There ure 5 testis, follicles and the vas 
deferens coils 6+ times around them. The 
spicules are stellate, 0,010.03 min in diumeter- 
Rentarkit P, paradoxun yar. mahenunt 
Michaelsen, 1920, from the Seychelles prob- 
ably represents a distinct species since it has 
only 24 coils of the vas deteretts and the 
spicules have 24 points in optical section. The 
shallow thoracic common cloacal system and 
the arrangement of the spicules beneath the 
superficial bladder cell layer is characteristic 
of the present form. Owing to the very shallow 
thoracic common cloacal space, the colony is 
especially firm. 


Didemaum candidum Savigny, Kott, 1972a; 19 
and synonymy, 

New Reeord; Elliston Bay, 

Records: see Kart 1972a. 
Deyeriptien: The present colonies have the 
usual dark-brown zooids with brownish-black 
Pigment cells, The surface test is thin but the 
basal test is slightly thicker. The cloacal sys- 
fem is thoracic although the primary canals 
may extend more deeply. The thorax of each 
goord is enclosed in its own test sheath as it 
crosses the common cloacal space. There ts 
sometimes a lateral organ near the posterior 
end of the thorax, There are 84 coils of the 
vas deferens around the single undivided testis 
tollicle. Dense spicules are present throughout. 
They are 0.03-0,05 mm in diam. and show the 
cheructenstic runge {rom burr-like to stellate, 
Vesicular cells previously described for Poly- 
syncraton orhiculam (sce Kott (9728) are 
present in circles around the branchial aper- 
ures. 


Remarks: The pigmented zovids and form of 
the cloacal cavity, and the single testis follicle 
with a large number of vas deferens coils 
around it, have been used to identify this 
species. The presence of the vesicular cells 
previously thought to be diagnostic of P. orbit 
enluemn 3s puzzling. However, although they 
havé not previously been described for 
Didemnnm candidum, they have previously 
been found randomly distributed over the sur- 
face of Didermniui moselevi (see Kott 1972a)_ 


Previews 


9 


Didenmom moseleyi (Herdman), Kolt, 1972a: 
19 and synenynyy. 
New Recores, Elliston Bay, Emu Bay (Kan- 
garoo. [.), Investigator Strait (Stas. X7, 
27, ¥21. Previous Records: sce Kott 1972a, 
Destription, The colonies are of the nsual Torm 
with dense white stellate spictiles, shullow 
thoracic common cloacal cavity, and large 
fateral organs. Living spectmens from Waterloo 
Bay are yellow-orange, encrusting red algae, 


Didemmum patulum (Herdman), Kott, 1972a: 
18 and synonytny-. 
New Record: Emu Bay (Kangaroo J,}, For 
Previous Records, Deseriptien, see Koti 
197 la, 
Renurks: The colony forms a faree investing 
sheet marked with tie usual blue grey lines Lo 
give a macbled appearance. 


Didemnum temstanum (Gottschuldt), Kote, 
1966: 287 and synonymy. Tokioka. 1967; 
77. 
Didemniodes 
[ROS8: 648, 
New Record: Elliston Bay, Previous 
Records: see Kaw 1966; Tokioka 1967, 
FIG. 36 
Description: The living colony is bright orange, 
although this is: lost in preservative, The spi- 
cules are small and spherical with many very 
short conical spines projecting from the sur- 
face, 0.02-0.03 mm in diam. and very dense 
throughout the test. The surface of the colony. 
is raised into mounds and ridges with coman 
cloacal apertures on the apex of tha thicken- 
ings. The primary ¢loacal canals extend almost 
jo the basal test and usually are posterior 
abdominal, while there are thoracic secondary 
canals, Zooids occur in large clumps anchored 
to the busal test by solid lest material in which 
the abdomina of the zoolds are embedded, 
while in the thoracic region there are separate 
test sheaths enclosing each zooid and con- 
tinuous with the surface test. The surface test 
is fairly thick, The mounds that are apparent 
on the surface of the test are therefore 
created by the great proliferation of zooids in 
these ateas, between the primary cloacal 
canals. There are 54 colls of the vas deferens 
wround a single Leslis Follicle, There is a long, 
oval lateral organ on each side of the thorax. 
The branchial siphon is fairly long with dis- 
tinet circular muscles 
Remarks: Although the common cloacal cavity 
in these specimens is not as well developed 


fernatarnunt Gouschaldr, 


180) 


us has been previously described for the 
species, the small spherical spicules help to 
distinguish it, Kott (§£972b) was not able 
ta identify # “yellow ecrustose” specimen of 
the family Didemnidae from West I. {near 
Penguin Rock), Although the gonads were 
not malire in the West I, specimen it has 
been compared with the present colonies from 
Elliston and is identicul in every respect, In 
particular, the cloacal system and the distrihu- 
lian and form of the spicules are identical, 


Leptoclinides reticulatus (Sluiter), Kott, 1972a: 
18 and synonymy. 
New Record: Investigator Strait (Stn, Y6). 
Previews Records: see Kotte 1972a. 


Description: Spindle and stellate, purple and 
orange, pigment cells are present in the surface 
layer of test. Common eloacal openings are 
frequently and evenly distributed over the sur- 
face. The common cloacal system is extensive 
at the aesophageal and posterior abdominal 
level, The zonids are of the usual form with a 
posteriorly directed altial siphon. 


Leptoclinides fungiformis n.sp. 

Type Locality: Outside Pearson 1, on 

gravelly bottom mostly attached to shell or 

rock fragments, 97,1969, 50 m, Shepherd. 

Raleyipe and Colype: SAM, E911. 

FIGS, 37, 38 

Description: Two specimens are avatlabte frum 
the type locality. They arc about 6 em high 
with a rounded head. There i @ single cloacil 
upetiure terminally, There is a thick dense 
layer of spicules in the surface test. at the level 
of the branchial siphons, Spicules are less 
dense elsewhere in the head. In the stalk. the 
spicules ure more evenly and densely distri- 
bated throughout. There is no suffuce layer of 
bladder cells. The test is firm but not tough. 
The test in the stalk is similar in consistency 
but is perforated by longitudinal spaces, The 
primary cloacal system consists of extensive 
cavities posterior to the superficial zaoid Jayer 
surrounding a central core of test. Secondary 
caitals receive the openings of the posteriorly 
directed strinl siphons and open into the pri- 
mary cloucul cavity. Zovids open to the sur- 
face of the colony by a 64obed aperture. The 
atrial apertures ate posteriorly directed from 
the posterior third of the thorax and hive dis- 
tinct circular muscle bands forming a 
sphincter. A circular sphincter muscle is 
present on the branchial siphon, but is pot quite 
so apparent There are also fine longluilinal 


PATRICIA KOTT 


muscles on (he thorax, There are 4 rows of 
aboul & sligmata on cach side of the thorax, 
Eges are lurge. The testis Follicle is apparently 
undivided and the vas défefehs is Wound 
around ft, The stomach is small and rounded. 
The zonids in these colonies are budding from 
the oesophugeul region, Embryos are present 
in a layer beneath the zooids, but none were 
sufficiently mature to discern their form. The 
posteriorly directed atrial siphons open into 
the secondary cloucal canis rather than 
directly into the common cloacal chamber, 


Remurks: Spicules are stellate and are fairly 
tures, Although closely related 1o Leprtoclinides 
Ringé in the development of the cloacal system, 
the present species is distingumshed by ils larger 
spicules und smaller zooids, by the single testis 
follicle and by the smaller intestinal loop. The 
present species is further distinguished by its 
weil developed stalk, No other stalked species 
of Leploclinides has previously been described. 


Suborder PHLEBOBRANCHIA 
Family ? 

Recerds; As well as the species of this <ub- 
order listed below, specimens of an un- 
described colonial species have been taken 
fron) stulions Y18, Y1I9 and from two vther 
Tocutions. all im Irivestigator Strait, The 
species will be described and its phylogeny 
discussed in a subsequent publicalion (Kott. 
in press (1)>, 


Family ASCIDJIDAE 
Ascidia thompsoni Kou, 19729; 27, 
New Recerd: Elliston Bay. 
Recverds: sce Kott 1972a- 
Peseriprion: Individuals are the usual aval, 
laterally flattened shape, lying mostly on the 
lefi side. The body is pinkish and shows 
throwgh the firm translacent testi, There are 
nO muscles on the left side of che bowky. Body 
musculature ix present as an trrégular network 
on ihe right or upper side of the bouy, stopping 
abruptly at the endoslyle, and no muscles were 
detecied on the left side of the body. There is 
not a row of short parallel bands around the 
ventral and dorsal border as in A. sydneyensiz, 
The peritubercular area is very shallow. The 
dersnl lamina is double in its anterior part and 
is ribbed on the left. side only, The branchial 
sic is simply folded between each tongitudinal 
vessel and there are simple papillae at the june- 
hon of the longitudinal and transverse vessels, 


Previous 


ASCIDIANS OF SOUTH AUSTRALIA IL 181 


Fig. 
Fig. 
Fig. 
Fig. 


Figs. 
Figs. 


Fig. 
Fig. 


Figs, 


Figs, 


33, 
34. 
35, 


36, 
37, 38. 
39-41. 


42. 
43. 
44-46, 


47, 48, 


Tridideninum cerebriforme, Spicules. 

Trididemnum spiculatim. Spicules. 

Polysyneraton magnilarvum, Colony. 

Didemnunr ternatanum, Spicules, 

Leptoclinides fungiformis, Pig. 37.—Calony. Fig. 38—Spicules. 


Metandrocarpa indica. Fig. 39.—Individual, Fig. 40—Mesial aspect of stomach. Fig. 41. 
—Later aspect of stomach. 


Symplegma arenosa. Individual. 
Stolonica azsiralis. Gut loop and stomach. 


Stolonica truncata, Fig, 44.—Iindividual. Fig. 45 —Body removed from test to show. body 
musculature. Fig. 46—Gul, gonads. 


Polyandrocarpa simulans. Fig. 47.—Calony- Fig. 48.—Gut, gonads, 


1&2 


Remarks; The present species it distinguished 
by the coudition of the dorsal lamina and the 
body musculature, 


Axcidia sydneyensis Stimpson: Kott, 1972a; 24 
and synonymy. 
New Records: Elliston Bay. Investigator 
Strait (Sin. ¥18). For Previour Records, 
Description, sec Kott 1972a, 


Order PLEUROGONA 
Suborder STOLIDOBRANCHIA 
Family STYELIDAE 
Subfamily povyzoinae 
Metandrencarpa indica n,sp, 


Type Leecation: Investiguter Serait (Stn. 
¥6), 23 m, scattered low recf{ with shell 
sund patches and strong surge, Watson, 
Helorype: NMV, HI59. Paratype: NMV, 
HIS8. 


FIGS. 39-41 
Descriptions The colonies cansist of sessile, 
round, laterally flattened, sandy Individuals 5 
mo m diam,, fixed to a common basal stalk 
which sometimes expands into a wider mem- 
hrane. The present colonies invest an algal 
stom. The zooids are close to one another but 
each ff entire and separate and the tests do 
not adhere. The test is thin and fairly brittle 
with embedded sand, Both apertures are an- 
terior and fairly close together. The branehial 
aperture is sessile and the atrial aperture is 
on a pointed siphon directed away from the 
hranchial aperture, The body wall is very 
delicate and closely adherent to che test. It has 
a fairly close mesh of very fine muscle fihres 
continuous all over the body. The branchial sac 
has 4 infernal longitudinal vessels on each side 
With 6-8 stigmata per mesh, crossed by paras- 
tizematic vessels, and there are 9 such rows af 
stigmata, The gut forms u simple open loop 
across the pesterior end of the body and the 
rectum ts produced anteriorly toward the base 
of the atrial siphon. The stomach is large and 
has about 14 conspicuous stomach folds. There 
is a longitudinal ridge along the lateral aspect 
of the stomach, which continucs to form a 
leng curved caecum im ihe gut loop, On the 
posterior side of this longitudinal ridge, the 
somach folds are parallel to it, "The folds an 
the anterior aspect of the stomach, however, 
extent! more obliquely and terminate against 
this caecal ridge so that they do not extend 
continuously from the pylone to the cardiac 


PATRICIA KOTT 


end af the stamach, “The anal border is smooth. 
The dorsal lamina ts plain. Unfortunately no 
gonads were detected, 

Remarks: Specics of the genus Metandrocarpu 
Michaelsen, 1922, have heen described with 
5-10 internal longitudinal vessels, und male 
and female gonads on both sides of the body. 
Although the present specics has only 4 
interhal longitudini#l vessels und the gonads 
ure nol developed, its stomach is similar to 
that found in M. dura (Ritter) and M 
michaelsen? (Ritter & Forsyth) both from 
California (sce Wan Name 1945), In both 
these species the stomach has a longitudinal 
ridge canlinuous with the pylori¢ caecum and 
against which the stomach folds terminate. In 
other gebera of Polyzoinue, eg., Allecorarpa, 
Theodorelia and Palyzoa, although there may 
sometimes be u small number of internal 
longitudinal vessels, the stomach is barrel- 
shaped with parsllel longitudinal folds. 


Symplegma arenosa n.sp. 
Type Locations Off Waldegrave [., Shepherd. 


Holoivpe: SAM, H904, Paratype: SAM 
EYRS. 
FIG. 42 
Description, The colonies are formed of 


sessile, sundy individuals, more or Jess pillar- 
shaped but laterally flattened, the zooids taller 
than their width. They are fatrly closely placed 
but are entirely separate and arise from a 
sandy basal plate formed of a tangle of basal 
stolons. Both apertures are close together on 
the upper free end of the body in a circular, 
sand-frec, area of verv thin test. The rest of 
the test is encrusted with sand, When the 
zooids are contracted, the stiffer, sand- 
encrusted, lest on either side of the apertures 
is drawn together across the openings which 
then appear to be depressed in a longitudinal 
slit on the supper surface. The body wall has 
strong muscles vround the anterior part of the 
body, consisting of longitudinal bands raciat- 
ing from both the siphons and short trans- 
verse bands extending across the mid line 
dorsal and ventral to both of the siphons, 
These short transverse bands (as in Agnesia 
eleciata; Kott 1969h) are instrumental in 
drawing together the protective test across the 
aperture. There are 4 internal longitudinal 
vessels on each side of the body and 15 rows 
Of stigmata, “Uhese are nol. crossed by paras- 
ligmatic vessels. There are 6-8 stigmata in 
each mesh, The longituding! muscles extend 
only about a third of the distanee down the 


ASCIDIANS OF SOUTH AUSTRALIA IL 


body, The gul torms a very short loop across 
the posterior end of the body and the rectum 
is long ind extends anteriorly toward the hase 
of the atrial siphon. The stomach is short and 
barreleshaped With about 14 longitudinal folds 
and there Is a short straight pyloric caecum 
connected to the anterior limb of the gut loop 
by a divided ligament and blood vessels (see 
Synpleenia oceania Tokioka, 1961), Vhere Is 
also i connective between the stomach snd ihe 


intestine. Unfortunately no gonads ate 
developed in the present specimens, 

Remarks: Although the present species 
resembles Metandrocarpa indica in the 


presence of 4 internal longitudinal vessels in 
the branchial sac und in the shape of the body, 


it differs In the body apertures, the body mus-. 


culalure, the shape of the stamach and pyloric 
caecum, arid in the length of the gut loop, In 
the present species, the stomach, as well as the 
number of internal longitudinal vessels, 
resemble the condition in the genus Syin- 
Plegnias and in Swnplegma vecania Tokioka, 
1961, from Noumea, the arrangement of the 
pyloric caecum and ifs ligaments and the con- 
nective between the stomach and the intestine 
are identical with the present species, Although 
Symplegni spp. are not usually upright as in 
the present species, they are joined (o basal 
stolons which form a mesh und eross one 
another and an upright form has been des- 
cribed once previously, viz: Syrmplegma viride 
stolonica Verrill (see Wan Name 1945), 

The condition of the apertures is. very 
specialised and the disposition of the body 
musculature to serve the protective device 
described above fas not previously been re- 
corded tn the family Styelidae, although similar 
mechanisms are developed in the families 
Agnesidae (see Kott 1969b) and Molgulidue 
(see Kot 1972b), 


Stolonica australis Michaelsen, Kott, 19724: 28 
and synonymy, 


New Record: Elliston Bay, Previous 
Records; see Katt }972a. 
FIG, 43 

Description; In colonies recently collected 


from Tipara Reef (Spencer Gulf), there is a 
single row of hermaphrodite gonads anteriorly 
on both sicdes of the body, but posteriorly there 
are tather irregular scattered series of simple 
testis follicles. There is some variation in the 
apparent shape of the stomach, from pyriform 
to barrel-shaped dependine on the condition 


3 


of the body wall cavenng the cardiac end, 
The caecum is only of moderate size and 
curved 


Stoloalea truncata f.sp. 
Type Location; | koe north-west nf Walde- 
grave Lon rocky bottom with sund patches, 
23 om, ILwI971, Shepherd. Helarype: 
SAM, E893. Paratyipess SAM E894, E909, 


FIGS, 44-46 
Description: Colonies comprise raunded, 
sandy, individuals joined by basal stoluus. Each 
individual js spherical, sessile, and about 1 cm 
in diam. The apertures are both anterior on 
the frec end of the body, and are present m 
conspicuous transverse interruptions to the 
father solid sand encrusted test, so that cach 
aperture is surrounded by delicate, thin test 
Which can be withdrawn and covered over by 
protective sand-strengthened lips, The body 
wall is Eairly muscular, with longitudinal bands 
radiating from both siphons, extending over 
the rest of the body, There is a broad band of 
circular fibres around the anterior end of the 
budy and associated with the protective clos- 
ing mechanism described shove. There ate 
also circulac fibres around dhe posterior end 
of the body, There are 3 branchial folds on the 


tight and 3 on the left, although they all tend 


to fade out posteriorly, especially the most 
dotsal folds, The internal longitudinal vessels 
are arranged according to the following 
formula: 
DL 0(10)5(9)5(6)6 E 

There are 14-2 stigmata per mesh, although 
these are set slightly obliquely in relation to 
the internal longitudinal vessels. The gut forms 
4 simple transverse loop with the rectum (lirn- 
ing slightly anteriorly. The stomach has a large 
number of very fine longitudinal folds, and the 
whole extent of the anterior border of the 
stomach is produced into a tube of peradually 
decreasing diameter, lined internally with glan- 
Jular folds in paralle] with those lining the rest 
of the stomach wall, This tube forms a curve 
in the gut loop and terminates in a pointed 
caecum, The anus is bilabiate, On the left hand 
side of the body, anterior to the put Joop, there 
may be 5-6 elongate and sometimes branching 
testis follicles, terminating distally in the long 
slender duct directed toward the atrial aper- 
ture. On the right hand side of the body, there 
are similar testis follicles arranged around the 
posterior half of the ventral border and around 
the posterior border of the body. Same of these 
testis follicles are also associated with an ovary 


184 


of 2-3 crys. lying on the mesial surface of 
the distal end of the testis and opening by a 
short wide oviduct, while the seminal duet 
curves uround and for most of ifs length lies 
free in the peribranchial cavity, There are up 
to 6 hermaphrodite gonads of this type in the. 
centre of the row of gonads on the right side 
of the body, Larvae are present in the right 
peribranchial cavity. They have uitudiate 
papillae andl appear to be of dhe type generally 
associated with the Polyzoinuc (see Millar 
1960), 


Remarks: The most unusual production of the 
anterior aspect of the stomach to form the long 
curved caccum: in the gut loop is quite dis- 
linctive, The long branching testis follicles are 
also unusual; while cxternally the highly 
specialised device to protect the apertures, 
each located in « transversely onented strip of 
unmodified! test, by withdrawing them beneath 
the firm sand-hardened test is unique. Closing 
mechanisms as described above for Svenple gira 
aréfosd tsp. and for specics in the families 
Molgulilae and Aynesiidae, usnally involve 
both apertures simultaneously. The test is black 
un can be seen through the encrusting sand. 


Ocolimuria australis Gray. Kott, 1972a: 29 and 
synonymy. 

New Record; Oll Waldegrave [, Elliston 

Bay, For Previors Recards, Description, see 
Kou (972g. 


Polvandrocarpa simulans n,sp. 


Type Location: Investigator Strait (Stn. 
Yo. ¥19), 23 m,.on scattered low-ceel with 
shell sand patches and strong surge, Watson. 
Further Reeordy: Anvestigntor Strait (Stn, 
Y19), Elliston Bay, St. Francis I. Holorype! 
NMY, HI62. Paratypes: NMY, TITAQ, 
H161, HL63. 
FIGS. 47, 48 
Deseriprion; The colonies form tight agere- 
gutes of Individuals about 1 cm in diam, The 
living specimens are reddish brown, tipped with 
black, although preserved specimens are black. 
The test is thick with 4 slight sand enerustation, 
although this as mul present internally where 
the test of adjacent zooids is confluent. Both 
uperlures ure sessile. The branchial aperture 
is terminal and the atrial aperture antero- 
dorsal. The body wall is very muscular, with 
ulmost continuous layers of tongitudinal and 
circolar muscle bands. The branchial sac has 
4 low fulus with crowded, very thick. internal 
longitudinal vessels. The transverse branchial 


PATRICIA KOTT 


vessels aire alsa thick so that the branchial sac 
is a Very tough organ in this species, Longi- 
twidinal vessels are arranged according to the 
following formula} 
E 1l6)1O9)1T(8IT(9I0 DL 

The body musculuture is especially thick 
around the apertires, The gut forms a small 
¢losed loop in the posterior end of the body 
and the rectum extends forward to the atrial 
aperture to form a secondary loop. A rout, 
Nt-tupped, enlocarp completely occupies the 
pomury gut loop as in Polycarpa pediunculate. 
The small, pear-shaped stomach has internal 
glandular folds. "The anal border has 4+ some- 
times indented rounded Johes. ‘The gonads are 
Nal, Nask-shaped ovaries opening by a short, 
wide, oviduct directed towards the trial aper- 
ture and overlying two tows of testis follicles 
with 5-6 follicles in each row. The ducts of 
each destis [ollicle join a common duet that 
runs along euch side of the avary to join 
logether and open into the peribranchial caviy 
on the mesial aspeet of the shurt oviduer. ‘Whe 
gonjds are embedded in thick body wall, There 
are up to S of these gonads on the Jeft side of 
the body, usually in a single Tow anterior fo 
the gut loop. On the right, there gre up to 9 
gonads usually arranged in 2 rather irregular 
rows, 
Remarks: This species is very reminiscent of 
Polycurpa pedunculata. In view ef the tight 
ugyregates and confluent test and the absence 
of sand between adjacent mdividuals. how- 
ever, il is clenr that colonics form by vegetative 
reproduction rather than by aggregation of a 
number of solitary individuals. The process af 
vegetative budding tu form these colonies is 
probably associated with the small size of the 
mature individuals, thus limiting the number of 
imternal longitudinal vessels in the brarchial 
sac and the number of gonads on the body 
wall, 


Polyandrocarps tapidosy = (Herdman). 
1952; 250. Millar, 1963: 730, 
Ciendyiric: lapidosa Herdman, 1899: 99. 

New Recered: Investigator Strait (Stn. Y19), 
Previews Records: Vio Port Phillip Heads, 
Westemport)—Millar 1963. NS.W, (Port 
Jackson) —Henlman 1899; Kott 1952. 

Deseriprian. The present specimen is a large, 

sandy. plate-like colony with the upper surface 

slightly concave. 9 em in diam, with a maxi- 
mum thickness of 2 cm, fixed by a large pan 
of the basal surface. The surface of the colony 
is smooth und sandy wathout conspicitous 


Kott, 


ASCIDIANS OF SOUTH AWSTRAITA Th 


swellings or furrows. The zooids are long, but 
both openings are on the upper surface, There 
are 4 long branchial folds of very varying 
height and internal Jongitudinal vessels are 
artanged according to the following fornmta: 
DL (10)1(3)1(7)1(4)1 EB 

The gut extends in a simple are from the 
postenor end of the body to the anterior atrial 
aperture, There are 12 internal longitudinal 
stomach folds. The gonads are elongate and 
arranged along either side of the endostyle. 


Subfamily BoTRYLUINAE 
Botrylloides Jeachi (Savigny}. Kott, 1972a: 29 
and synonymy. 
New Records: Elliston Bay, off Waldesrave 
[. For Previous Records, Descriptian, sec 
Kon 1972a, 


Botrylloides magnicoecum Jtarimeyer, Kott, 
1972a; 30 and synonymy. 
New Records: Waldegrave 1,, Pearson I. For 
Previous Records. Deseriprion see Katt 
1972, 


Rotryllus schlosseri (Pallas). Kotl, 19724: 31 
and synonymy. 
New Record: North of Waldegrave [, For 
Previous Records, Description, see Kott 
197 2a, 


Subfamily STYFLINAE 
Styela plicata (Lesueur), Kott, 1972b: 239 and 
synonymy. 
Ascidia plicate Lesweur, 1823: 5. 
New Record: Coflin Bay. For Previous 
Reéeords, Description, sec Katt 1972b, 


Styela pedata (Herdman). 
Polycarpa pédata Herdman, -188l: Tl, 
Tokioka, 1958; 322 and synonymy, Kott, 
1964; 137, 
Pardocia pedata Hartmeyer, 
1360, 
Stvela whitelegit Kott, 1952: 213. 
Telhyuent whitelegii Hartmever, 1909-112 
1364. 
New Recordy: Off Waldearave I,, Pearson I, 
Preyious Records: N.S.W, {Port lackson, 
Port Stephens, Port Cnrtis)—Herdman 
1899; Kott 1952. Qld. (Moreton Bay, Great 
Barrier Reet|—Hastings 1931; Kott 1964, 
Indonesia—Pizon 1908. Phitlipines—Herd- 


1909-11: 


184 


man 188l: Van Name 1913. 

Tokioka 1958, 

FIGS. 49, 30 

Description: The preserved specimen is orange, 
the test ix tough and leathery, with longitudinal 
ridges without any foreign bodies attached. 
The body is of characteristic shape. expanded 
postero-dorsally, Both apertures: are directed 
upwards and the branchial aperture, on a 
siphon continuous with the upright ventral 
surface, is often recurved, The atrial aperiure 
iv also on a short siphon above the postero- 
dorsal expansion of the body. Basally the test 
Is extended into prop-like roots. The body wall 
is muscular. The dorsal tubercle iy oval with 
numerous separate circular openings giving it 
®& porous, sponge-like appearance. 

Up to 21 internal longitudinal vessels are 
evenly spaced on the folds, and 3-7 internal 
longitudinal vessels are present between the 
folds. There are 6-8 stigmata per mesh, The 
wut forms a fairly wide loop with tall endo- 
carps enclosed by the loop. The stomach 3s 
ciliptical with longitudinal internal glandular 
folds, The anal border is lohed. 


There are 3 beanched and ramifving gonads 
on the left ubove the gut loop und up to 2 
on the right. The gonads are embedded in and 
eccupy most of the body wall cxcept where 
the gut is present on the Jeft. Tall endocarps 
are present over the body wail between the 
branches of the gonad. The yvonads have 
testis follicles closely applicd to either side of 
the ovary. 


Remarks: The gonads and endocarps of this 
species are very similar to the much branched 
and ramifying gonads of S. ramiificata Kott, 
1952, which has also heen recorded from 
Moreton Bay. S. ramificara is, however, a very 
much smaller species heavily encrusted with 
sand and shell, with a continuous V-shaped 
Opening on the dorsal tubercle, a narrow put 
logp and a longer rectum than in the present 
species. A similar dorsal tubercle is present in 
Polyearpa aurata (Quoy & Gaimard) (see Van 
Name 1918) which is also similar to the 
present sptcics im other characters. It is dis- 
tinguished mainly by its short, typically poly- 
carp gonads in contrast to the long ramifying 
stvelid gonads of the present species, Sryele 
pedata, therefore, has a wide distribution from 
the Phillipines and Japan and around the 
custern seaboard of Australia to the Great 
Australian Bight. It overlaps with P. ayrerfe in 
the Phillipines and on the east coast of Aus- 
tralia. 


Japan— 


186 


Polycarpa tinctor (Quoy & Gaiimard), Kutt, 

1964: 134 and synonymy. 

Ascidia tinctor Quoy & Guimard, 1834: 

608. 

New Record: Off Waldegrave T. Previouy 
Records: see Kott 1964, 

Description, A single specimen only is avail- 
able. tt is large and shyhtly damaged, The test, 
however, has the usual hard, brittle, sand- 
encrusted lorm characteristic of the species, 
The specimen tg laterally flattened, with the 
wiritl aperture hall way down the dorsum, The 
apertures are sessile. The dorsal tubercle is 
Jarge with a complementary slit, The branchial 
sac has 4 very narrow folds. Gonads were not 
detected in the present specimen, 
Remarks: The present specics had not pre- 
viously been taken further south than Port 
Jackson. It does occur commonly on the north- 
West and north-east Australian coast, in the 
East Indies, and off Japan. The form of the 
dorsul tubercular slil, broken into several 
parts, has previously been described in speci- 
mens from Japan and. the East Indics. 


Polycarpa pedunculata Heller. Kott, 1972a: 35 
and synonymy, 

New Records: Blliston, N-W of Waldegrave 

I., 22 m off Waldegrave L, Pearson 1., Inves- 

tigator Strait (Stns. X9, 15, 19, 21, 25, 27; 

YG, 12, 23; Z9, 11), Emu Bay {Kangaroo 

L.). For Previous Records, Description, see 

Kott 1972a. 

FIG, 51 

Remarks; Specimens in this collection demon- 
strate the full range in external uppearance, 
from sessile to stalked individuals, from brown- 
ish to black individuals, In general, the stalk 
of specimens froni Investigator Strait is longer 
than that found mn specimens from ‘St, Vincent 
Gulf, and many of the specimens are super- 
ficially very similar to specimens of Pyure 
scoresbiensis with which they occur, demoan- 
strating convergence in their external appear- 
ance relalecl to the environment, 


Polycarpa clavata Hurlmeyer, Kott, 1972a; 33 
und synonymy. 
New Retwray: Walldegrave 7, 22 m off 
Waldezrive L, Pearson T., Investigator 
Strait (Sta. Y¥21). For Previous Records, 
Description, see Kott 19728, 


Family PYURIDAB 
Pyura spinifera (Quoy & Gaitiatd), Michael- 
sen, 1922: 390 (part: Port Jackson speci- 
tens}, Katt, 1952: 269; J972a: 39. 


PATRICIA KOTT 


Ascidia xspinifera Quoy & Gaimard, 1834: 
617. 

Boltenia australiensis Carter, (885; 197, 
Bolrenia niberculata Herdman, 189: 511; 


1899; 17, 
Cynthia multiradicara Herdman, 1899: 
30. 


Bolrenia spinifera Michaelsen, 1YO5! 72 

(part: not B, gihbosa) , 

Bolrenia spinesa var. tntermnedia Michael- 

sen, 1908: 390, 

Pyura glbbasa var. intermedia Michelsen, 

1922; 390, 

Pyura australiensis 7. tvpica Michaelsen 

& Hurtmeyer, 1928; 419, 
New Records: Off Waldescave b., Investi- 
gator Strait (Stns, YU9, Z?). Previous 
Recards: W. Aust. {Albunyi—Quoy & 
Guimard 1834. S. Aust. (St. Vincent Gulf) 
—Kott [972a, Vie. (Port Phillip Heats, 
Portland Harbour, Cape Woolami, Bass 


Strait)—Carter 1835; Michaclsen 1995. 
NSW. (Mort Jucksonj}—Herdman 1899; 
Kott 1952. 


FIG, 52 


Remarks: Minute scalelike spines are present 
on localised ridges in the branchial siphon. The 
branched keratose fibres (see Carter 1885) 
and Michaelsen & Hartmeyer 1928) are pre- 
sent in the body wall and stalk. They are 
similar to spicules found in Pyura stolorifera 
(see Millar 1962), The specimens are invar- 
iably covered with a sponge (Halisarca; sec 
Carter 1885). 


Pyura australis (Quoy & Gaimard), 

Pyurq australis subspecies. auxtraliv Kel, 

1972a: 39 and synonymy. 

Pyura «australis var, parvispinalis Kott, 

1952; 26%. 

Baltenia gibbasa Herdman, 1899: 19. 
New Records: Off Waldegrave 1, NW. of 
Waldegrave I., St. Francis I, Pearson I., 
Investigator Strait (Stux. X27, ¥21), Emu 
Bay (Kangaroo 1.). Por Prevlous Records, 
Descripiion, see Kott 1972. 

FIG, 53 
Remarks: The principal character on which the 
distinction between the two subspecies, P. aius- 
tralis typica and P. australiy purviypinatis, was 
based is: the condition of the anal border. A 
careful ¢xamination has shown that in this 
group of species Lhe lerminal part of the rec- 
tal wall always has flat folds projecting into the 


ASCIDIANS OF SOUTH AUSTRALIA 


lumen as describe for Po australis parvise 
pindtiy Kott, (952. These folds are usually 
extended beyond the anal border mto rounded 
Jabes which are often Jong and finger like and 
sometimes subdivided, Occasionally. however, 
they do not extend oitside the anal tim. There 
is no constant condition observed for any 
group of specimens in regard to the anal hor- 
der and it is concluded that subspecies ara wot 
indicated. The slight difference in the length of 
the branchial spines ¢(Kott 1952) is ot signifi- 
cant, These are conical and only slightly curved 
and arise from a long oval hase. 0.02-0.05 
min Jong, and their length along the spine ts 
0,020,04 mm. The largest spines are hearest 
to the upertures, Typical stellate spicules, 0.02 
mm in dium, wre always present in the test 
und siphonal lining and distinguish the species. 


Pyora pachyderinatina (Herdman) $,sp, draschii 
nom, nov, 
Boltenia pachvdermating Herdman, 1881; 
81. Drasche, | 884: 370. Herdman, 1899: 
16. 7Herdman & Riddell, 1913: 875. 
Pyura pachydernmutrina var, gibbosa Kott. 
1952: 265. 
New Records: Waldegrave L. Elliston Bay. 
Previows Records: W. Aust (Cottesloe to 
Albany}, Vic, (Julia Percy 1., Flinders, 
Walkerville)—Kott 1952. NISW. (Kiuma, 
Port Jackson )—Drasche 1884; further speci- 
mens in Austedliun Museum: 
FIGS, 54, 55 
Bescripion: Typical specimens with dumb-bell 
shaped spicules and fringed anal lobes, The 
siphonal spies are produeed into a long 
pointed hase which js distinct from the rounded 
base in P. australis, or the flattened scaletike 
base in P. xpinifera, Michaelsen (1905, 1908, 
1922) and Michaelsen & Hartmeyer (1928) 
utiempted rationalisation of the relationships of 
the spegics Pye pachydermatina, P. gibbose 
and P. spinifera and their synonyms, on the 
basis of external appearance, condition of the 
Salk and of the dorsal tubercle, and the 
presence or absence of spicules. A study of 
the group in Australia has shown that the form 
of the spicules and siphonal spines, supported, 
withio certain Jumits, by the form of the anal 
border, provides. the only reliable character on 
which to distinguish the species, Michelsen 
& Harnmeyer (1928) characterise FP. pachy- 
dermatina by the presence of a smooth anal 
border, complicated dorsal tubercle, and dumb- 
bell shaped spicules. This is anly true, how- 
ever, for the New Zealand subspecies FP. 


187 


pachydermatina typica since although the 
duorb-bell shaped spicules ure wlways present, 
the Australian forms never have a smooth anal 
border. The dorsal tubercle may be a Jess com- 
plicated double spiral cone. Thus Pyara gib- 
hose; Michaelsen & Hartmeyer 1928 (type 
speciinen: Cynthia sibbosa Heller, 1878, from 
Bass Strait) with anal Jobes and dumb-bell 
shaped spicules falls within the definition of 
P. pachydermatind and Heller's specimen be- 
comes the type of the subspecies P. pactiveder- 
nuninde epibbosa (net P. pechydermartine var. 
gibhosa Kott, 1952), Pyura gibbose intermedia 
Michaelsen, 1922 (>P spinifera intermedia 
Michaelsen, 1905) trom Backstairs Passuye, S, 
Aust, together with P. spinifera typica (part: 
P. tuberculaia Herdman) from New South 
Wales, neither of which have dumb-bell shaped 
nor stellate spicules, anc consequently syn- 
onyms of P. spiniferd, rather than af P. pacliy- 
dermatina gibbova, Pyura  pachydermatina 
draschii is distinguished from P, pachiyder- 
matine gihbosa (>P; pachydermutine inrer- 
viedia; Katt 1952) (which has shallow anal 
lobes) by its fringed anal border. P. Pachyeer- 
marina gibhasa overlaps the present subspecies 
from Bass Strait to Port Jackson, and extends 
Further to the north. The western limit for P. 
pachydermatina draschit is not at Walkerville, 
Victoria, as Kott (1952) had suggested, since 
it extends to Western Austialta. 


Pyura scoreshiensis Kott 1972a: 36, 
New Records: Pearson L, Investigator Strait 
(Sens. MM it. 15, 17, VIA) Previous 
Records; see Kott |972a, 
Description; Specimens available show a wide 
range in the length of stalk and in the develop- 
ment of the rooting processes at the base of 
the stulk. These sometimes form large sandy 
lamellue as the roots spread and sand adheres 
to them, The apertures are always close 
together on the upper surface, although the 
pronounced ridge between the apertures. pre- 
viously descrbed (Kott 1972a) is not always 
present, 
Pyora irregularis (Herdman), Kott, 1972a: 
38 and synonyrny. 
New Reeord: Investigator Strait (Sen. V6) 
For Previous Recerds, Description. see Kott 
1972a, 


Pyora tendata asp. 
Type Lucation: ltwvestigator Strait (Stn, 
¥21), 30 m, scattered, low, on pebble reef 


1X8 


Figs. 
Fig. 
Fig. 
Fig. 
Figs, 


Figs, 


Fig. 
Fig. 


49-50. 
51. 
52, 
53. 
34, 55, 


56, 57, 


38. 
59. 


PATRICIA KOTT 


We Danie 


Styela pedata. Fig. 49.—Individual. Fig. 50—Gonads from left side of body. 

Polycarpa peduncalata, Gonad showing flask shaped ovary and testis follicles. 

Pyura spinifera, Fig. 52a—Branching spicules from body wall. Fig. 52b—Branchial 
spines, 

Pyura australis. Spicule and branchial spines. 

Pyura pachydermarina. Fig, 54.—Dumbell shaped spicule and branchial spines. Fig_,55. 
—Anal border showing internal folds from rectal wall. 

Pyura tendata, Fig. 56.—Longitudinal section to show sandy coaling around body.-—Fig,. 
57—Body opetied around ventral surface, branchial sac removed to show branchial 
velum, tentacles, dorsal ganglion, gut and gonads: branchial sac (bs); body muscles 
(bm); branchial velum (v), branchial tentacles (bt); dorsal tubercle (dt); dorsal gang- 
lion (dg); dorsal lamina (dl); intestine (i); gonad (g); base of the atrial siphon (as); 
anal opening (a). 

Herdmania momus. Individual. 

Malgula ellistoni, Gonads, gut loop and kidney on inner surface of body wall. 


ASCIDIANS OF SOUTH AUSTRALIA II 


with sand patches, J. Wain. Holotype: 
NMV, H1L56, 


FIGS, 56, 57 

Desevipitons The species appears to occur in 
aggregates. The holotype, however, is the only 
complete specimen available and only small 
parts of at least two others were obtained when 
the specimen was broken [rom its substrate, 
The outline of the individual iy not obvious 
superficially, since jt js completely surrounded 
by a sandy cout 1 cm or more thick. The test 
is verv thin, without wrinkles, and minute 
hair-like extensions of the test are scen extend- 
ing across a narrow space belween the sandy 
coating and the test, It is probably these hairs 
which enmesh the sand forming the thick coal 
ing around the body, The narrow space 
between the outer surface of ihe test and the 
sandy coating is occupied by various commen- 
sal worms and echinoderms, 

The animal itself consists of a club shaped 
hody narrowing to a long terminal branchial 
siphon, The atrial siphon fs twice the length 
of the branchial siphon and extends forwards 
from the dorsal border of the body in the 
anterior part of the posterior third of the body 
length and opens level with the branchial open- 
ing, Excluding the sandy coat, the body is 1 
cm deep from its dorsal border at the base of 
the atrial siphon to its ventral border. The 
length of the atrial siphon is 2 cm, Owing to 
the gradual narrowing of the body to the base 
of the branchial siphon, the body appears to 
have two long diverging siphons of equal 
length, 

The body wall is very muscular, Branchial 
tentacles are present at the base of the 
branchial siphon half way between the exter- 
nal aperture and the hijse of the atrial siphon. 
Just anterior to the ring of branchial tentacles, 
which are 3 times branched, there is a large 
muscular velum protuding into the lumen of 
the branchial siphon. The dorsal tubercle is 
in the base of a very long warrow peritither- 
cular area extending half way down the 
branch! sac, Tt is u very small, simple UL 
The clongate dorsal ganglion is associated! with 
the base of the atrial siphon just posterior to 
the dorsal tubercle. There are 9 branchial folds 
with internal longitudinal vessels arranged as 
follows: 

E (4)0(7)0013)1(15)2(1.6)3(2)1016)2013 07) DL 

The gut forms a simple loop in the posterior 
end of the body. There are minute, branched, 
liver tubules in the pyloric region and the 
body wall covering the distal portion of the 


189 


Tectum is produced into a pronounced non- 
muscular arial velum so that the bilabiate anus 
actually opens into the base of the wtrial siphon 
beyond this velum, The branchial velum bas 
very strong circular muscles in its basal hall, 
The body musculature i very strong, with 
outer circular bands forming an almost con- 
tinuous coal external to strong well-spaced 
longitudinal muscle bands. The gonads consist 
af about 7 or 8 pairs of polycarpike sacs 
vither side of central ducts in the gut loop on 
the left amd in a corresponding position on the 
right side of the body, 

Remarks: This quite extraordinarily modified 
species is in most essential aspects typleally 
pyurid and resembles Pyara cancellaia Brewin 
(see Kott 1971) in the thick sandy coating 
created by sand enmeshed by the lony test 
hairs all around the bacdy, Its most conspicuous 
character is the very long atrial siphon and the 
numowinyg of the anterior part of the hody 
so that the branchial aperture is also pro- 
duced upwards to u level with the atrial siphon, 
so that the incurrent ciliary slream is not oh- 
structed by the sand being accumulated around 
the body, The thick, sandy coating is so dense 
and so rigid that it is hard to imagine how the 
animal is able to increase in size. It is probable, 
however, that the commensals present between 
the sandy coating and the test constantly irtt- 
gate this ares and thus maintain the spice into 
which the animal can expand as it grows 
These commensals are therefore probably 
essential to such sand covered species. (see 
also Pyvra cancellata; Kot 1971). 


Halocynthia hispida (Herdman). Kotte 1968; 76 
and synonymy; 19772a; 4]. 
New Record: Investigator Strajl (Stn. X19). 
aoe Previous Records, Description, see Kot 
1963_ 


Herduiania momus (Savigny). Koti, 1972a; 
41 and synonymy. 
New Records: Off Waldegrave I., St, Fran- 
cis T,, Pearson L, Investigator Strait (Sto. 
X17). Spencer Gulf, For Previouy Records, 
Description, see Katt 19724. 
Reewrks: There is the usual great range In size 
ot individuals, Smaller specimens are mere or 
less upnght with transparent test and short 
furrowed siphons. Larger specimens become 
faccid and opaque. 
Ctenicella antipoda Kott, 1972a: 44, 
New Record; Investigator Strait {lubcl 
Hlegble). For Previews Recards, Dexerip- 
tien, see Kott 19724, 


7490 


Remarks: The distal part of the gut loop is 
distended with mud. In ong large specimen the 
fohuds on the left side of the body are enclosed 
in the gut loop. In view of the agreement with 
the type specimen in all other aspects, this 
must be regarded as. an individual variation. 


Mokguta eflistoni n.sp. 


Type Loedion: Elliston Bay, in caves out- 
side bar, subject ta strong swell, 14.v.L971. 
Holowpe and Purai'pest SAM, E907. 


FIG, 59 

Description: Small sandy spherical individuals 
were found adhering to Evlerdmania ausrreis 
und sometimes forming aggregates. Both aper- 
tures are present fairly close together on the 
upper suttace and are directed away from one 
another. VYery delicate muscles radiate out 
trom the siphons. There are internal circular 
muscles around the branchial and atrial 
siphons. ‘There aro 7 branchial folds on each 
side of the body, with f meshes along each 
fold. Each mesh contains a primary spiral in- 
Fundibulum, which subdivides into two half 
way up into the fold. here are 3 internal 
longitudinal vessels arranged along the fold, 
There is the usual long gut loop open at the 
pole and a short curved kidney on the right side 
of the body. The gonads consist of a cireular 
ovary with a short, wide, duct directed dor- 
sally. Testis follicles are present along the 
proximal border of the ovary and join into a 
very shore vas déferens which opens into the 
penbranchial cavity on the mesial surface of 
the ovary. ‘the right gonad is in the middle of 
the hudy wall and the left gonad is in the 
secondary gut loop. The specimens are abour 
)}S cm in diam, 


Renwrks: The position add form of the gonads 
with the very short vas deferens opening on the 
surface of the ovary are distinctive. Anural 
embrycs qre present in the peribranchial cavity, 


Molgula subolosa Quoy & Gaimard 1834: 613. 
Koll, 1972b: 248 ancl synonymy. 


New Revord: Elliston Bay, For Previous 
Records, Deyerintian, see Kotl 1972b. 


CharucterisGes of the Faun 


Afi outstanding feature of the fauna is the 
large number (23) of species in the relatively 
primitive Aplousobranchis. Polycieridae, 
Clayvelininge und Euherdmaniimae, in which 


PATRICIA KOTT 


common cloucal systems ive not developed. A 
further 7 species ure secondarily colonial 
species of Styelidue; one is 4 colonial phicbo- 
branch species: three ure aggregated pyurids: 
and there are two aggregated spccivy of the 
family Molgulidae. Thus about half uf the 
species present, although colonial in habit. pre- 
serve theit own independent. openings and do 
not form cloacal systems. They clo, however, 
demonstrate rematkuble morphological adap- 
tations in the arrangement, orientation and 
operation of them apertures ta maximise their 
Teactions. with the environment. Colonies of 
Ritterella herdmania, Plucentela ellisien’ n-sp., 
and the new colonial phlebobranch species are 
of special interest in that the “Zooids ure 
arranged in parallel, so that their branchial 
and atrial apertures are respectively loeatecl on 
different sides or at different ends of the 
ealony. Some benefit can accordingly be 
derived from mutual reinforcement of feeding 
currents. One would expeet that the exact 
erientalion of these colonies would be such 
that prevailing currents could also reinforce 
the incurrent ani excurrent ciliary streams, 
These species and many of the Clavelininac. 
together with the new species Merandraecgrpa 
indice, Syinplegma arenesa and Stolenica trim- 
cata, all have specially adapted body muscu- 
Jature to operate sophisticated siphonal uppara- 
tus, Adaptation ol the nervous system. to serve 
the specialised musculature can algo be 
expected, In fact, im both Pyura fendata and 
in the new colonial phlebobranch species, the 
feucal gland anid ganglion ure localed pos- 
teriorly in association with an unusual 
posterior position of the alnal siphon. 


None of the species discussed ubove ure 
enefusling ant most favour habitats under 
ledges and in crevices or caves. There are 
anly 18 species with highly evolved cloues! 
systems that have jichieved a degree of inde- 
pendence of the enviranment and, with only 4 
eavenlions (Didemanm cendidum, D. moselesi, 
Trididemnum spiculatuoc and Bowryllus schloy- 
sere), these jire jilso butky or stalked (rather 
than enerusting) and favour habitats under 
ledges or in crevices or caves. 


Phe majority of the solitury stolidobranch 
specics that are present are stalked and able 
fo maximise their seactions with the environ- 
ment hy moving on their stalk with the current. 
There are only few solitary botton-living 
species (Asetdia spp., Polycarpa  rnetor, 
Crenicella antipoda) or large fixed species 
(Merdmiania mamus, Styela pedata), 


ASCIDIANS OF SOUTH AUSTRALIA II 


yi39 


GHENT AUST HALL AN FLAT 


ize 3350 1? 


SOUTH AUS RAL i 


Fig, 60. Map of South Australian locations from which specimens are recorded. 


Zoogeography 

The known ranges of Atapozoa marshi and 
Pseudadistoma australis have heen extended 
from Western Australia to the eastern end of 
the Great Australian Bight where they overlap 
with Distaplia distomoides, Euherdmania aus- 
tralis, Pseudadistoma cereum and Polyandro- 
carpa laupidosa, whose ranges have been simi- 
larly extended {tom the easi. The ranges of 
Aplidium rubricellun: and Palyclinum nep- 
tunium from the west, and Ascidie thampsoni, 
Pyura irregularis and P. stolonifera (see Kott 
1952) from the east. are also known to extend 
into either Spencer Gulf or St. Vincent Gulf. 
Thus the distribution of these ascidian species 
across the south coast supports the existence of 
a marine faunal boundary at the eastern end 
of the Great Australian Bight, separating the 
Flinderstan marine faunal Province from the 
Maugean Province to the east (Knox 1963). 
There is less eyiderice however for a western 
boundary of the Flindersian Province further 
south than Cockburn Sound in Western Aus- 


tralia. It is possible that the 15 new species 
described herein will subsequently be found to 
have wider ranges across the Australian coast 
und provide further data relating tao the 
western boundary of the Flindersian Province 


‘to help resolve the question of the existence of 


a Western Australian Province (Baudinian of 
Kott, 1952; see Knox 1963). 


The ascidian fauna of Gulf waters of South 
Australia includes only a limited number of 
the species that occur along the open coast. 
Of the species discussed. above which terminate 
their range off the eastern South Australian 
coast, only Aplidium rubricollum, Polyelinum 
neptunium, Aycidia thompsoni, Pyura irregu- 
laris and P, stolonifera extend into guilt waters, 
Podoclavelia. meridionalis, Distuplia ytylifera, 
Didemnum ternaranum, Styela pedata, Poly- 
carpa tinctor, and Pyura pachydermutina are 
also apparently absent from gulf waters 
although their geographic range around the 
Australian coast is wide, It is probable, there- 
fore, that some ecological factor inhibits the 


bye 


spread ot the ascidian fauna Erom the open 
const Inte gulf waters. It is possibly the same 
factor Usar favours the occurrence af Distaplig 
viridis, Aplidium pliciferum, Bideninum lani- 
bitin, B. psendodiploxoma, Leptoclinides rufus, 
f, kKingt, Eehinaclinum verrilli, Asetdia aclara, 
Palvearpa papillata, Pyura cataphracta and 
Microcosmus nichollsi in St. Vincent Gulf and 
other eimbayments to the east and nerth around 
the Australian coast, although they have not 
been recorded from the open coast. The ascidian 
fauna of gulf waters of South Australia is 
therefore distinet from that of the adjucent 
cogstal waters and has no special zoogeo- 
graphic affimty with either the Flindersian or 
the Maugean Provinces. 

There are & further 42 species discussed 
herein that are recurded from both coastal and 
eulf waters. The majority of these have a 
Wide cusmupulitan distnbution or an extended 


PATRICIA KOTT 


range from Western Australia to cither the 
houndary of the Maugean marine faunal revion 
in Bass Strait (Hedley 1904, Knox. 1963). or 
further up the castern Australia coast. Also 
included in this group of species, however, are 
Crenicella antipada, Pyura scoresbiensixs and 
Atapezoa fcititeustana, previously thought en- 
demic t St. Vincent and Spencer Gulfs, and 
Aplidiuin colelloidey (see Kot 1972a)- 


Acknowledgements 

Grateful thanks are due lu Mrs. }) Watson, 
who collected the material from Investigator 
Strait for the Nutional Museum uf Victoria, 
and Mr. S$. A. Shepherd. of the South Austra- 
ian Department of Fisheries and Fauna Con- 
servation, who made the collections from mest 
other locations. 


‘The work was done while the author was the 
recipient of A.R-G.C. Grant D65/ 15386. 


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Appendix I—Station List 


Stations are listed in the order north to south and 
west to east. 


DENTAL BAY. (Coll. I. Thomas). Lat. 32°13’S; 
Long, 133°38’E, 
Near Ceduna; in Posidonia community; depth 
10m; 1965. 
Mfapuzoa fantayiana 
Sycacoa cérebrifermix 


OFF ST. FRANCIS 1. (Coll. 8. Shepherd), Lat. 
32°31'S: Long. 133°15'R. 
Rocky bottom; slight current; depth 55 m; 
8.1971, 
Pyendodisiorma cereum 
Palyandrocarpa simulans 
Pyura australis 
Herdmania momus 


WALDEGRAVE ISL. BND. eo $. Shepherd). 
Lat, 33°36'S; Long. 134°46 
Strong surge; depth 20 m; a iv.1970, 

Distaplia distomoides 

Synaicinm papilliferum 

Botrylloides magnicoccum 

Batryllus schlosseri (22 m) 

Pyura australis 
Rocky bottom; 
23.x.1970, 

Enxherdenania australis 

Aplidium colelloides 

Oculinaria australis 

Botrylloides leachi 

Styela pedate 

Palycarpa tinctor 

Pynra australis 

Pyura spinifera (covered with sponge) 

Pyura puchydermatina 

Herdmania momius 


strong surge; depth 22 m; 


1 Km N.W. OF WALDEGRAVE T, (ANXTOUS 
BAY). Lat. 33°33'S; Long. 134°46'E. 
Rocky bottom; slow current; depth 23 m; 
11.¥,1971, 
Glaveling mirahiliy (attached to limestone). 
Distaplia distamaides 
Distaplia stylifera 
Syinplegsma arenosa 
Batryllus schlosseri 
Sloleonica truncata 
Holozoinae sp. (grawmg on red algae) 
Polycarpa clavata {growing on brown or red 
algae: Sargassum or Osmundaria) 


Herdmania montis (with red algae ajtached) 
Rocky bottom with sand patches; slow current; 
depth 23 m: 11.v.197). 

Podoclavella cylindrica 

Synoiciunt papilliferum 

Stolonica truncata 

Polycarpa pedunculata 

Pyara australis 
Racky bottom; 
23.x. 1970, 

Cluvelina nodyla 

Polyeitor giganieum 

Pyendodistoma anstraliy 

Psendodistoma cereum 

Aplidium flavalineatymn 

Symplegnia arenosa 

Polycarpa clavata (attached to rock) 

Palycarpa pedunculata 


slight surge, depth 22. m; 


WATERLOO BAY (ELLISTON). (Coll, S. Shep- 
herd), Lat. 33°38'S; Long. 134°51'E. 
Roof of caves; strong surge; depth 6 mi; 
13.v.1971. 
Clavelina bhaudinensis 
Padoclavella molaccensis 
Distaplia distomoides 
Endistoma renieri 
Pseudodistama cereum 
Aplidinen lobatium 
Polysyneraton paradoxtm 
Didenripn candidum 
Didemnum maselevi 
Ayctdia thampsoni 
Asctdia sydueventsis 
Polvandrocurpa sitiulans (growing on Ascidia 
avdueyensiv) 
Chitside bar; very strong surge; depth 17 mm 
12v.1971. 
Sycozaa cérebrifarmis 
Polvelinum nepluninm 
Aplitinm elatum 
Ocnulinaria australis 
Pyura pachydermazina 
Outside bar, in caves: strong swell, depth 17 
m; 14.y.1971,. 
Pyenoclavella diminuta 
Fuherdmania rnustralis 
Ritferella herdmania 
Placentela ellistont 
Trididemnaum cerebrifornte 
diminita) 
Stolonica australis (around base of FEuherd- 
mama australis) 
Molgula ellistoni (adhering to E. Australis) 


(investing P. 


ASCIDIANS OF SOUTH AUSTRALIA II 15 


Inside bar: depth 6 m; 12.v,1971, 
Palycitor giganteum 
Psendodistoma australis 
Didemnum moseleyt 
Palycarpa pedurrcilata 
Inside bar; strong surge; on vertical face; depth 
6G m: 12.v.1971, 
Aplidium pantiterinim 
Near entrance to bay, in caves; depth 3 m:; 
l4.v. 1971, 
Pseudodistona exrnusense 
Palyelinam neptuniaum 
Motlenta sabulosa 
Floor of cave; depth 16 mz: 12.v.1971. 
Atapozoa mirahilis 
Centre of bay; depth 11 m; 12.V,1971. 
Atapezoa mirabilis 
On root of caves: depth 3-5 m: 14.v.1971, 
Aplidium flavalineatum 
Flliston Bay 
Cystidyles dellechiajei 
Trididemnum spiculatum 
Didermmaum ternatanam 
Oculinaria australis (with EB. 
M. ellisteni) 
Ratryllaides leachi 


australix and 


PEARSON ISLAND. (Coll. S. Shepherd). Lat. 
33°56'S; Long. 134°15’F, 

Rough-water coast, 400 m offshore on gravelly 
battom, attached to shell or rock fragments; 
moderate surge; depth 30 m: 97.1969. 

Leptoclinides fungiformis 

Péalycarpa clavata 

Pyura australis 

Herdniania mamus 
Rough-water coast; in caves; moderate surge; 
depth 35 m; 101.1969. 

Poadoclavella meridionalis 

Aplidium rubricoltum 

Stvela pedata 

Polveurpa peduncelata 

Herdmania momus 
Sandy beitom hetween Dorothee and Veteran 
Is, Lat. 34°1I'S; Long, 134°15’E. Slight surge; 
depth 70 mz 11.1,1969. 

Palycitor giganieum 

Batrylloides magnicoecum 

Polxcarpa pedunewlata 

Pyurg scoreshiensis 
Outside Pearson 1; depth 35 m, 

Padoaclavella meridionalis 

Leptoclinides faungiformiy 

Styela pedata 


COFFIN BAY. (Coll. 8. Shepherd). Lat. 34°38'S; 
Long. 135°30' F. 
Oyster Trays, at low water; slow currents; no 
sediments; October 1979. 
Sivela plicata 


SPENCER GULF. (Coll. S. Shepherd). Lat. 
34°2'S; Long. 137°23'E. ‘Tipara Reef: depth 11 
m; 20.vii.1971., 

Pycnecluvella diminwiv 

Herdmania mamus 


INVESTIGATOR STRAIT (Coll. J. Watson). 
Station X7: depth 30 m: 101,1971. Lat. 
35°16’S; Long, 137°30'E. 
Didemnunt moseleyi 

Station X9; depth 31 m; 19.j.1971. Lat. 35°17°S; 

Long. 137°30'R. 
Palycarpa pedunculata 
Pyura scoresbiensis 

Station X11, depth 30 m: 

35°19" S; Long. 137°30°R. 
Pyura scoresbiensis 

Stalion X15; depth 32 m; 

35°23'S: Long. 137°30’E. 
Polyeitar gigantelm 
Polysyncraton magnilarvim 
Polyearpa pedunculata 
Pyura scaresbiensis 

Station X?, 19.1.1971. 
Palvandrocarpa simuyltans 

Station X17; depth 35 m; 

35°24'S; Long. 137°30'E. 
Trididemaum cerebrifarme 
Pyura scoreshiensiy 
Herdmiania momuy 

Station X19; depth 34 m; 

35°26’S; Long. 137°30'F. 
Ruherdmania australis 
Colonial stolidobranch—Gen. 
Axcidia s¥dnevensis 
Polycarpa pedunculata 
Haloeynthia hispida 

Station X21: depth 34 m: 

35°28’S; Long. 137°29'E. 
Polycitor giganteum 
Polycarpa peduncufata 


Station X25; depth 35 tn; 

35°31'S; Long. 137°29'R, 
Polycarpa peduncnlata 

Station X27; depth 31 m: 

35°33'S; Long. 137°29'F. 
Didemaum moseleyi 
Palycarpa pedunculata 
Pyura australis 

Station Y6: depth 23 my; 28.1.1971. Lat. 35°17'S; 

Long. 137°16'E, 
Leptoclinides reticulatits 
Polysyncraion magnilarvum 
Metandrocarpa indica 
Polyandrecarpa simulans 
Polyvarpa peduncilata 
Pynra irregularis 

Station Y12; depth 33 m; 

35° 23'S; Long. 137°17"E. 
Palycarpa pedunculate 

Station Y14: depth 32 m; 

$5°25'S:, Long, 137°17’E, 
Alapozou marshi 
Pyura scoreshicusis 

Staton Yil6; depth 35 m: 

35°26'S; Long. 137°17'E. 
Sycozea peduncalata 

Statton Y17; depth 34 mm; 

35°27’S8; Long. 137°18'E. 
Atapozoa marshi 

Station Y18; depth 31° m; 

35°28'S; long. 137°18'E, 
Oxycorynia arenosa 
Polycitor obelixcaum 
Colonial stolidobranch—Gen. & sp.7 
Ascidia sydneyensis 


T9i1971, Tat. 


WDG1971. Lat. 


19.1,1971. Lat, 


20.4;1971, Lat. 


and sp.? 


17.i,1971, Lat. 


175,197], Lat, 


17.41.1971. Lat. 


24.0.1971. Tat 


233.1971, Lat, 


DLIO7L. Lat. 


205.1971. Lat. 


23.1,1971, Lat. 


196 


Station Y19; depth 33 m; 20.1.1971. 
35°29'S; Longe. 137°18'E. 


Lat, 


PATRICIA KOTT 


Station 2? 
Colonial stolidobranch—Gen. & sp.? 


Polyandrocarpa lapidosa 
Polyandrocarpa simulans 
Pyura spinifera 


Station ¥21; depth 32 m 


35°32'S: Long. 137°18'E. 
Aplidium colelloides 
Polyedrpa clavata 
Pyura australis 
Pytra tendata 


Station Y23: depih 32 m; 


35°33'S: Tong. 137°18'E. 
Polycarpa pedunculata 

Station Y?; 207.1971. 
Didemnum niaseleyt 


201.1971. Lat. 


17.4,1971, Lat, 


Pyura spinifera 


Stations Z9, 11; depth 38 m; 25.11.1971. Lat. 


35°30'S; Long, 137°8’E. 
Polycarpa pedunculata 
? (Lahel illegible) 


Colonial stolidobranch—Gen, & sp.? 


Ctenicella grtipoda 


KANGAROO ISLAND, (Coll. I. FE. Watson). 
Lat. 35°35’S; Long. 137°31'E. Off Emu Bay, 


221,197], 
Didemnum moaseleyt 
Didemnum patulum 
Polycarpa pedunculata 
Pyura australis 


Index to Genera and Species 


Page 
Aplidium colelloides 176 
Aplidium elatum nsp. _ - 177 
Aplidium flavolineatum 176 
Aplidium fobatum ,, : 176 
Aplidium pantherinum .. 176 
Aplidigm rubricollum 176 
Ascidia sydneyensis 182 
Ascidia thompsoni .,,, 180 
Atapozoa fantasiana 168 
Atapozoa marshi : 168 
Atapozoa mirabilis n.sp. 168 
Botrylloides leachi - Sete 185 
Botrylloides magnicoecum __ 185 
Botryllus schlosseri 185 
Clavelina baudinensis ,, . 166 
Clavelina mirabilis n.sp. 165 
Clavelina nodula n.sp. .... T66 
Ctenicella antipoda 189 
Cystodytes dellechiajei ..., 172 
Didemnum candidum ..., 179 
Didemium. moseleyi 179 
Didemnium patulum 179 
Didemnum ternatanum 179 
Distaplia distomoides 170 
Distaplia stylifera ..,, 170 
Budistoma renieri _ - : 171 
Euherdmania australis ... 172 
Halocynthia hispida 189 
Herdmania momus cant 189 
Leptoclinides fungiformis n.sp. 180 
Leptoclinides reticulatus 180 
Metandrocarpa indica n.sp. ..., 182 
Molgula ellistoni n.sp. _ 190 
Molgula sahulosa . 190 
Oculinatia australis 184 
Oxycorynia arenosa nsp. 167 


Page 
Placentela ellistoni a,sp. 173 
Podoclavella cylindrica 167 
Podoclavella meridionalis 167 
Podoclavella moluccensis 167 
Polyandrocarpa lapidosa 184 
Polyandrocarpa simulans n.sp. 184 
Polycurpa clavala - - 186 
Polycarpa pedunculata ..,, 186 
Polycarpa tinctor 186 
Polycitor giganteum 171 
Polycitor obeliscum n.sp. 171 
Polyclinum neptunium . 175 
Polysyncratan magnifarvum _ (78 
Polysyneraton paradoxum .., 178 
Pseudodistoma australis (72 
Pscudodistoma cereum 173 
Pseudodistoma cyrnusense 173 
Pycnoclavella diminuta 170 
Pyurg australis 186 
Pyura itregularis _.. 187 
Pyura pachydermatina 187 
Pyura scoresbiensis 187 
Pyuri spinifera 186 
Pynra tendata n.sp. 187 
Ritterella herdmania 172 
Stolonica australis 183 
Stolonica truncata n.sp, 183 
Styela pedata , 185 
Styela plicata. ., : oF 185 
Sycozoa cerebriformis .... 170 
Sycozoa pedunculata 170 
Symplegma arenosa n,sp, 182 
Synoicium papilliferum .  _ 177 
Trididemnum cerebriforme ..., 178 
Trididemnum spiculatum 178 


STRIGEATA (TREMATODA) OF AUSTRALIA BIRDS AND MAMMALS 
FROM THE HELMINTHOLOGICAL COLLECTION OF THE UNIVERSITY 
OF ADELAIDE 


BY GEORGES DUBOIS AND L. MADELINE ANGEL 


Summary 


An important collection of Strigeata of birds and mammals has been made by the Department of 
Zoology of the University of Adelaide. 

The present work records thirty species, of which seven are new. Descriptions of the new species 
with additional information on some of the others are given. The new species from birds are: 
Apatemon (Apatemon) vitelliresiduus (from Biziura lobata), Cardiocephaloides ovicorpus (from 
Phalacrocorax melanoleucos brevirostris and P. varius), Cotylurus (Cotylurus) magniacetubulus 
(from Cygnus atratus), Diplostomum (Diplostomum) parvulum (from Hydroprogne caspia and 
Pelecanus conspicillatus), Neodiplostomum (Neodiplostomum) lanceolaturn (from Ninox 
novaeseelandiue). 

The new species from mammals are: Neodiplostomum (Triloborchidiplostomum) diaboli (a new 
subgenus, for which the diagnosis is given) (from Sarcophilus harrisii) and Pharyngostomoides 
dasyuri (from Dasyurus viverrinus). 


STRIGEATA (TREMATODA) OF AUSTRALIAN BIRDS AND MAMMALS 
FROM THE HELMINTHOLOGICAL COLLECTION OF THE UNIVERSITY 
OF ADELAIDE 


by Georces Dusots* and L. MaveLine ANGEL] 


Summary 


An important collection of Strigeata of birds and mammals has. been made by the Department of 
Zouloxy of the University of Adelaide. 

The present work records thirty species, of which seven are new. Descriptions of the new species 
with additional information on some of the others are given, 

The new species from birds are: Apateron (Apatemon) vitelliresidues (from Bittura lobar), 
Cardiavephaloides avicoraus Grom Phalacrocerax melanoleuces. brevirestris and P. varias), Cotyluras 
(Corylarus) magniacetubulus (from Cyenns atratus), Diplostomum (Diplostomum) parvalum (from 
Hydroprogne caspia and Pelecanus conspicillatus), Neodiplostomum (Neodiplostomum) lancealatum 
{from Ninax navaeseélandiue ). 

The new species from mammals are: Nerdiplostamum (Triloborchidiplostemum) diaboli (a new 
subgenus, for which the diagnosis is given) (from Sarcephilus harrisii) and Pharyngostomoides dasyuri 
(from Dasyvurus viverrintes). 


Resume 


Une importunte collection de Sirigeala d°Otseaux et de Mummifares a été constituée au Deéparte- 
ment de Zoolagie de Université d’Adelaide, 

Le présent travail comprend Ja description ou ta mention de 30 espéces, dant 7 sont nouvelles: 
Apateman (Apaternen) vitelliresiduus, Cardiocephaloides ovicorpus, Cotylurus (Cotylurus) magniaceta- 
Inilus, Diplostomum (Diplostomum) parvelum, Neodiplastomum (Neadiplostomtm) lanceolatum, Ned- 
diplostamum (Trilehorchidiplostamum) diaboli (n. subgen., dom la diagnose est proposée) et Pharyn- 
rostumoides dasyuri. Ces deux dernigres espéces sont parasites de Marsupiaux (Dasyurinés), respec- 


tivemenot de Sarcophilus harrixil et de Dasyurus viverrinus. 


Introduction 


An important collection of Strigeata from 
birds and mammals has been made by the De- 
partment of Zoology of the University of Ade- 
laide. It consisted of 92 tubes with spint 
specimens, and 3 slides, 

Collections and identifications made before 
August, 1951, are the valuable contribution of 
the late Professor T. Hurvey Johnston, to 
whost memory this work is dedicated. Since 
that time Mrs. P. M. Thomas (Patricia M. 
Mawson) has done most of the collecting. 
One of us (L.M.A.) collected the remainder. 
Dr. J, C. Pearson, of the University of Queens- 
land, Brisbane, contributed 5 specimens 
(Pharyngesromoides dasyuri) from Tasmania. 

Some of the specimens are valueless, either 
because of poor preservation or because young 
stages cahnot be identified with adults. 

‘The holotypes of all the new species des- 
cribed in this paper have been deposited in 
the South Anstralian Museum (SAM), Para- 
types, where »vailahle, are in the South Aus- 
tralian Museum, and in the Helminthotlogical 


~ . Collection of the Institute of Zoology, Univer> 


sity of Neuchatel (G.D_). Preparations of the 
rest of the material are deposited in the Uni- 
versities of Adelaide and of Neuchatel. 

The present publication is a continuation of 
3 previous accounts of Australian $Strigeida 
(Dubois. 1937; Dubois & Pearson 1964, 1967). 

Thirty species are described or recorded. 
Seven, one of which belongs to a new sub- 
genus. are new. Twenty-seven are recorded as 
u¥ian parasites anc the other three are from 
Taammals. 


Family STRIGEIDAE Riuillict 
Subfumily STRIGEINAE Railliet 


Apharyngostrigen simplex (S. J. Johnston, 
1904). Dubois, 1968: 35, figs, 19-21. 
Dubois & Pearson, 1965: 79, figs, 1-3, 
S. J. Johnston, 1904: 112, pl. 7, figs. 1-3- 

Host and origin; Ardea novachollandiac 
Latham, from Tailem Bend, River Murray, 
5. Aust., 1.v.1940 and 9.xi1,1940 (2 specj- 
tens); from the Australian Museum, date? 
{one specimen). 


*Tustitut de Zoologic, Université de Neuch&tel, Suisse, 
| Department of Zoolozy, University of Adelaide, Adelaide, S Aust. 5000 


Trans. R. Sor. S, Aust. Vol. 96, Part 4, 30 November 1972 


198 GEORGES DUBOIS and L- MADELINE ANGEI 


Hahitat: upper intestine. 
Deseription® These specinens, taken from. the 
type-host, measure 2,8-3.0 mm; eggs 92-99 
by 61-68 em. 
Parastrigea, repens (Chase, 1924), Dubois, 
1968; 68, fig, 51, Chase, 1Y21; 500, fig. 1 
and pl. 26 figs. 1-5. 


FIGS, |, 2 

flast and wviging Circuy approximans Peile, 

from Tailem Bend, 8. Anst.. 1Qav. 1950 (18 

adult and 1 immature specimen) ind Dee. 

1938 (10 young specimens), 

Neahitat; duodenum. 

Chase (1921) found three specimens of this 
strigetd in the intestine of Neropheyx novere- 
hollanediae (Lathany) Srom Terrigal, N.S.W. 
The holotype, which is registered in the Aus- 
tralian Museum (W544), was reexamined by 
Mr, J, C. Pearson (see Dubois 1968, p68, foot- 
note 1). On the basis of this examination, the: 
species owas removed from the  yenus 
A phuryngostrigea Ciurea tu Perastrigea Sziduat. 
Nescription: The smallest specimens: with few 
eges in the uteri measure 1,6-2,2 mm. Fully 
mature worms are 5-6 mm long. Suckers 
weakly developed: oral sucker marginal, 60- 
115 pm in average diameter, ventral sucker 92- 
165 yan, near oral sucker, Average ratio of 
the oral to the ventral sucker nearly 2? =: 3. 
Length of forebody Sram 9-19 faverage 14) 
times that of oral sucker, Protealytic gland 
elongated, oval or fusiform, 190-220 by sO- 
110 ,»m, composed of closely averegated 
lobules and lying between the two cencentra- 
tions of the vitelline follicles, 

Ovary kidncy-shaped and testes multi-lobed, 
occupying second hall of the hindbody. Vitel- 
Jariy of forcbody extending dorsally up to 
ventral sucker, concentruted in Jateral semi- 
cordifarm expansions of dorsal lip of tribocy- 
tic organ. (No follicles tn ventral lip, which is 
as long as forehody), Scattercd follicles in wall 
of segment, extending further forward dorsally 
than ventro-laterally; In hindbody, virclline 
follicles concentrated in front of ovary, absent 
dorsally Over the gonads, and extending ventro- 
laterally to the bursa copulatrix, Bjaculatury 
duct joming with alerus at entrance to genital 
conc. Eggs numerous. 90-105 hy 60-68 am. 
Reluionships: Parastvigea repens, P. inrer- 
inedta! Tubangui, 1932, and P. flexitis (Dubois, 
1954) are closely related apharyngeal strigcids; 


in the uhsence of pharynx they are distinguish- 
able from all known members of the genus, 
Their normal hosis are birds of prey {Falconi- 
formes). P. iniermecdia (feam the Philippines) 
differs from P. repens in that the small suckers 
are subequal, and in the size of the eggs (100 

112 by 71-79 nm), PF. flexilis is distinguished 
from che Australian species by having fewer 
ceys, and by «a geographical distribution 
festeicted Lo the holaretic zone of Europe and 
Asia 


Parastrigea sp, 
Host and origin: Uhreskiornix moluece 
(Cuvier), from Queensland, 26vi.191L (4 
contracted speciinens. collected hy A. 
Breinl), 
Rauhite: unknown. 


Desenipiion! Body fength |.4-2.4 mm-. Fore- 
body 0.88-1.08 by 0.94-1.15 mm (dorso-ven- 
tral diam.), with two well-developed lateral 
expansions. Hindhody 0.75—1.40 by 0.81--1,05 
mm. Bursa copulatrix small. 

‘These worms exhibit a striking similarity to 
BR. robusta Szidat. 

A specific diagnosis is reserved until better 
specimens are obtained. 
Strigea baylisi Dubois. 1937; 1968: 82. figs. 

6-61, 

Hoss and orivin: Theeskiornis meolucea 

(Cuvier), from Tailem Bend, 8, Aust, 

28.41.1942. (one specimen), Platelea flavipes 

Gould, from Toilem Bend, 24.17,1943 (23 

specimens) and 10.x11.1947 (6 specimens). 

Habirar. intestine: 
Relationships: Strigea baylixi, which appears 
to be @ parasite of Platalcidac, is distinguish- 
uble from S. promiscua Nicoll by its smotler 
size, the smallness and the weakness of the 
phurynx (30-60 by 37-47 um), the extension 
of the vitellaria to nearly as far as the posterior 
extremity, and the absence of follicles from the 
ventral wall of the anterior scgment, 


Strigea glandulosa Dubois, 1947: 244. fic 9; 
1968) JOL, figs. 82-84. Dubois & Pearson, 
1965; 82, figs. 4—5. 

Strigea falconixy Dubois, 1937: 247, fig. 
10 (not Szidat, 1928). 
lavix and origin: Circus approxinums Peale, 
from Tailem Bend, 8. Aust., 15.xii 1938 (2 
young specimens), MValiastur  spherurus 
(Vicillot), from Tailem Bed, Dec. 1958. (1 


One of us (G.D-}, having examined three syalypes of P. intermedia, has not found any trace of 7 


pharynx, 


STRIGEATA OF BIRDS AND MAMMALS 199 


specimen} and 18.¥i,1941 42 macerated 

specimens). Faloo submiger Gray from Men- 

ingic. S. Aust. 6.v.1945 (4 specimens y- 

Habitat: intestine. 

Deseripliian; Body ip to 2.8 mm in length 
(when extended). The smallest specimen, 
with eggs in the uterus. sacasures only 1.4 mm, 
Oral sucker terminal. often prominent, 120— 
140 by 90-125 ,m, pharynx rounded. very 
muscular, 95-105 hy 90-105 jm; ventral 
sucker 160-190 by 150-175 ,m: proteolytic 
gland well developed, oval, strongly lobulated, 
145-210 by 170-250 yim, lying wt base of fore- 
body, 

Ovary reniform. Mehtis’ gland intertesticular. 
well developed. Fggs 89-102 by 57-69 ym, 
veTy NUMETOUS i Muture specimens, in which 
the uterus may he distended into sinuotis or 
lortugus curves and even into Joops, 
Relavonships: Sirigea plaidulose differs Erom 
5. foleenis Szidat in its small size and in the 
greal development of the proigalytic pland 
relative lo the length: of the body, 


Strigea nicolli (Dubois, 1937), Dubois, 1968: 
114, figs, 101-102. 
Sirigea sutiont Dubois, 1937: 237, figs. 5— 
7. 
Hass and origin: Gyninorhina  ihicen 
fLatham) from Canberra, A.C.T., April 
1969 (4 specimens) and 27.vii,1960 £1 
specimen; collected by R. Mykytowyez), 
Flebirar: duodenum and intestine. 
Descriptian: Length 1.40-1.62 mm. Oral 
sucker 121-12 by 195-177 pm: pharynx §5- 
LOS by 75-90 ym; ventral sucker 190-230 by 
215-230 pm, Eggs 108-115 by 65-72 jm. 
Relarionships: Strigea nicolli resembles S. bay- 
fis! Dubois in general anatomy, but differs in 
the sizes of the suckers and the pharynx, which 
are definitely Jarger, and in the minor develop- 
ment of the atrial ring-shaped musculature. 


Srigea promiscua Nicoll, 1914: 347, Dubois, 
1968; 119. figs. 107-1083. Dubois & Pear- 
son, L967: 1R6, 

fHlaxts and origins Ninox jevaextelandiac 
(Gmelin} (syn. N. boofeek) from Yatkuri. 
S. Aust. 24.viil. 1987 (1 specimen, deserihed 
below). N, strenaa (Gould) fram Bidsvold, 
Qid,, 1vil9L9 (22 specimens, very con- 
tracted; collected by M. J. Bancroft). 
Habiraty intestine, 
On June 9, 1965, Dr. I. C, Pearson collected 
the species from the smal} intestine of Ninox 
Hovaeseelandiae jn Brisbane, Qld, The species 


has now been found four times, always from 
the same host genus. 

Description; Length 2 mm, Body very con- 
tractilo, nearly as wide as long when. strongly 
contracted (as in 1,¥7,1919 material), Oral 
sucker 160 by 185 ym, pharynx 115 by 105 
ym, ventral sucker 230 by 210 ym; proteoly- 
tic gland 190 by 215 ,m, multilobed. 

Ovary renitorm, 170 by 260 «mm. Testes 
roughly lobed. the anterior measuring 285 by 
390 ym, the posterior 320 by 400 xm. Vitel- 
lara extending from cephalic margin to level 
of equator of genital conc. The latter is robust, 
well differentiated. larger than the ovary, 260 
zm in diam. when retracted, Genital atrium 
spacious, L80 to 240 jum in depth, with ring- 
shaped musculature well developed. Eggs (04— 
115 hy 75-80 pm, 6 in number. 

AS at present known, it seems that Sinigea 
premiscud 1 restricted 4o the Singiformes. 
especially to the arientul type Nivea 


Apatemon (Apatemon) vitelliresiduus n.sp- 
FIG, 3 

Host and origin: Biziura lobata (Shaw) trom 

Tavem Bend. S. Aust., 10.x11.1937 and 

9.x. 1940 (35 specimens); Sandgate, Qld. 

22.18.1918 (1 specimen); Pumong and 

Caloot, R, Murray, S. Aust, 20, vi l958 (10 

specimens, obtained from two hosts) (type 

material). 

Habitare intestine. 

Holotype: length 2.6 mm, SAM, E927 [with 

3 paratypes. E928, on same sliile). 
Descriptien: Body cambered dorsally, 2.2=2.4 
mm long, Forebody ovoid or cup-shaped, 0.71-. 
0.90 mm in Jength by 0,51-0.64 mm in dorso- 
ventral diam., delimited by constriction from 
hindbody. Hindbody twice as long, banana- 
shaped, with small bursal region slightly de- 
lineated, 1.48-1.70 mm long by 0.50-0.56 mm 
in diam, at the Jevel of the tesies, Ratio of 
hindbady to forebody ringing from 1.9-2.1, 
Oral sucker 115-160 by 110-127 «am, terminal 
in position: pharynx smaller, 85-93 by 70-75 
pmMe ventral) sucker postequatorial, relatively 
large, 190-210 by 165-180 um, Proteolytic 
gland small, rounded and lobed, situated dor- 
sally near base ef forcbody, 63-80 by 60-70 
pM, 

Ovary 140-150 wm Jong, 180-190 am thick. 
located at anterior ihird (27-36/100, average 
33/100) of hindbody, Testes latge, upproxi- 
mately equal in size (first, 260-330 by 240- 
300 wm, second 300-370 by 260-300 gm), 
ovoid, roughly lobed, obliquely orientated with 


GEORGES DUBOIS and L. MADELINE ANGEL 


a4 


=o 
ae 


STRIGEATA OF BIRDS AND MAMMALS 


lobes directed forward. Vitellaria are confined 
ventrally in hindbody,, especially well developed 
in front of gonads, extending to near posterior 
extremity of Worm, but not masking bursa 
copulalrix laterally; erratic follicles extend 
more or Jess far into ventral wall of forebody, 
but do pot go beyond acetabular level, Vitelline 
reservoir lies in intertesticular space. Mebhlis” 
Slabd is situated dorsally and a litle anteriorly. 
Genital cone of mediam size, 190-200 by 150— 
180 nm, not well differentiated, Vesicula sem- 
inalis S-shaped and voluminous, lying postero- 
dorsal to second testis. Sinuoux ejaculatory 
duct opens into terminal part of uterus at 
entrance lo conc. to form a rectilinear, not 
pleated, hermaphroditic canal. Eggs 87-95 by 
58-67 pm, average 92 by 63 pi. 
Relationships: This new species closely 
resembles Aparemon (.4.) fuliguiae Yamaguti, 
1933, in geeral morphology, but differs essen- 
tially in the presence of erratic vitelline follicles 
in the ventral wall of the anterior segment, This 
femnant appears to constitute an archaic 
character. In 4, fitliguiae. the eggs are larger 
(100-220 by 60-66 um). 


Apatemon (Australapatemon) — intermedius 
{$. J. Johnston, 1904), S. J. Johnston, 
1904: 109, pl. v, figs. 7-10. Dubois, 1968: 
169. figs. 162-164. Dubois & Pearson, 
1965: 85, figs, 6-7. Johnston & Angel, 
195i; 66, figs, 1-28, 

Hosis and origin: Oxyure australis Gould, 
from Tailem Bend, S. Aust., 15.1.1941 and 
28.1941 (11 specimens), Accipiter fas- 
ciatus Vigors & Horsfield from Mallala, 8. 
Aust., March [965 (20 small ovigerous 
specimens). (fohnston & Angel (1951) 
found this species in 5 of 11 Cygnus atrartr 
(Latham) from Tailem Bend). 

Habitat: unknown. 

Description: Body Jength 1.5-2.8 mm (speci- 

mens from Oxyurer wustraliz), This species is 

characterised by the structure and size of the 
genital cone; this organ measures 240-320 by 

160-200 ,m when retracted, thus being ahaut 

one fifth the total length of worm, A wide 

strongly folded hermaphrodite duct passes 
through the cone. Laurer's canal descends from 


20) 


oviduct where the latter leaves the ovary, and 
Teaches the dorsal surface on a level with an- 
terior leslis, Eges 94-99 by 60-68 am. 

The specimens from <Accipiter fasciatus 
{possibly an abnormal host) measure 1.2 to 
1,9 mm. Eggs 90-95 by 55-63 pm, 


Curdiocephaloides hilli (S. J. Johnstan, 1904), 
Dubois, 1968; 180, figs. 175, 176; 1970: 
722. S. L Johnston, 1904; 110, pl VIL 
figs. 1-3, 


FIGS. 4, 6 

Hest and origin: Larus novaehollandiae 

Stephens, from Glenelg. 8, Aust., 2.iii.1939 

(20 specimens, from 2 hosts}; West L, 5. 

Aust. I4.vL1968 (4 specimens}; St. Kilda, 

S. Aust., 5-19.ix.1951 (2 specimens, from 

intesting! residucs of four birds). 

FHatbirar: duodenum, intestine. 

The only. previous record of this strigeid is 

that of §, J. Johnston (1904) who described 
it from the Australian gull. Lartx novaehollan- 
diae from Jervis Bay, N.S.\W. There is only 
one syntype (deposited in the Helminthological 
Collection of the London School of Hygiene 
and Tropical Medicine (No. 2447) known, 
The following description is based on the 
examination of 15 specimens, slightly smaller 
than those of Johnston, from the Glenelg 
material, 
Description: Body length 3.5-6.2 mm (6.9- 
§.2 mm according to Johnston}. Forebody 
ovoid in lateral view, subcordiform in ventral 
view, with feebly developed lateral expansions, 
®.74-1.12 mm tong, 0.55-0.70 mm. thick. 
Hindbody elongated, cylindrical and usually 
flexed dorsally, with bursal region set off from 
Temiuinder by a more or less definite constric- 
tion, 2.80-3.18 mm long. 0.52-0,80 mm in 
diam, at level of testes, 0.65-0.76 mm in hur- 
sal zone. Ratio of hindbody to forebody rang- 
ing from 2.9-3.8, averaging 3.4. Oral sucker 
terminal or subterminal, of medtum size, 95— 
LT? by 85-104 jm; pharynx smaller, spherical, 
RO-B5 by 75-95 ym, caeca extending back 
close to genital cone (fig. 6); ventral sucker, 
95-130 by 118-145 pm, lying usually just in 
front of middle of forebody_ 


—_ ee esesesssssssssssSse 


1, 2. Parastrigea repens, from Circus approximans. Fig. t—Lenath 5.5 mm, flattened mature 


Apatemon |Apatemon) vitelliréesiduns asp. fram Biziure lobata. Holotype: length 2.6 


Figs, 
m specimen. Fig. 2.—Length 1.8 mm, young specimen with two eEBS 
Fiz. 3 
mm. 
Fig, 4. Cardiocephalnides bill, from Laens novae-hallandiae. Length 3,64 mm, 
Fig. §, 


Univ, (G.D.) No. V17. 


Cardtocepleatoiites muxculnsas, from Hydreprogne caspiqa. Outline. Length 6.6 tant. Neuch. 


202 


Ovary, 13-151) hy [90-200 yim, situated at. 
about mid-léength of hindbody, in front of 
reliitively smiull testes [TAfE 190) by 210-250 
pm), whiele ure ovoid and roughty lobed. 
Vitoria confinedt to this segment, where they 
uppew profuscly developed up to gonads, 
then restricled to ventro-lateral field; they doa 
NOL penetrate bursa copulatrix. Viteiline reser- 
voir lies in intertesticular Space. WDilatahle 
hursa copulatrix large, quite often larger than 
forebouly; occupying last third or quarter of 
hindbody, aad enclosing powerfully built geni- 
tal cone, which measures 520-740 by 340-8001 
am; inner wall ol conc. when retracted, thrown 
into a number of folds delimiting sinuaus 
spaces. Uterus and muscular ejaculatery duct, 
the walls af which are S-10 ,m thick, enter 
the cone, proeced side by side and unite to 
form avery sherl hermaphraditic canal which 
discharges into large genital acrium. There are 
two sphincters, one ut orifice of cone, the 
other surrounding aperture of bursa. The 
Timerous eggs mensure 115-125 by 73-84 um, 
averuye 119 by 80 jm; shell thin (3-4 pm)- 


Cardlocephaloides muscolosus (S. J, Johnston, 
Tad), S.J, Jahnston, 1904; 112, pl VE, 
figs. +—9. Dubois. L968: 188, fig. 185. 


FIGS. 5, 7 

Hoste and erisin: Chlidonias hyhbrida 

(Pallas), from Tailem Bend. $. Aust. 

27.%.1948 (6 specimens). Aydroprogne 

caspia (Pallas), from Townsville, Old., 

20.vili. 1968 (1 Specimen). 

Habitat: small intestine. 

The only Australian record of this. strigeid 
is that of S. J. Johnston (1904), who described 
it rom the crested tern, Slernu bergil Lichten- 
sicin, from Broken Bay, NS.W. The type 
material collected by Dr. J. P. Hill cannot be 
found, As a result of the examination of these 
new specimens, some udditional structures have 
been seen, 


Description: Body up to 6.8 mm. long. Fore- 
body short and curdifonm, or pear-shaped 
(seeq side view), with large lateral expansions 
in last two thirds, 1.43-1.57 mm long, 1,60- 


GEORGES DUBOIS und L. MADELINDE ANGELI 


1.65 mm wide, 0,69-0,87 mm thick across 
the cephalic cupule, 095-115 mar at level 
ef expinsions, Hindbody clongated, subcylin- 
drical and slightly flexed dorsally, gradually in- 
creasing in diqm. Lowards posterier end where 
it is truncated, expecially just in front of bursa 
copulatrix, 445.2 mm tong, 0,72-0.87 mm 
in diam, anteriorty, 1.05-1.25 mm posteriorly, 
Rawio of hindhody to foreboily from 3—3.5. 
Oral sucker spherical, of medium size, measur- 
ing 130-160) ym in diam pharynx smaller, 
125 pm; ventral sucker. 130-140 pm, tying 
much in Front of middle of forebody {about 
at one-tnird its length), where latter begins 
ta dilate, Caeca extending laterally to Jevel of 
genital utcium, 

The ovary and the testes—the latter lobed 
und approximately equal in size (300-350 p.m 
long, and wider than long)—occupy about the 
middle third er third yuaster of the hindbody. 
Vitellaria cunfined to hindbady, profusely de- 
veloped in small follicles, obscuring the con- 
tents, duwn to the gonads, then restricted to 
two ventro-fateral fields. and extending dewn 
to gap of the genital cone, laterally beyond 
ity e@yuutor. Vitelline reservoir between testes. 
Large bursa cepulatrix «a truncated cone, 
occupying a little Jess than last quarter of 
hindbody, delincated by a slight constriction, 
gradually increasing in diam., with wide open 
terminal aperture; enclosing a powerfully built 
ovoid genital cone. 1250-1600 by 950-1160 
pm, clearly defined by its own musculature. 
When withdrawn, its walls are thrown into 
number of folds surrounded by parenchyma 
and muscle fibres. and delimiting sinuous 
spaces. A slack sphincter surrounds orifice of 
conc. Uterus entering cone anteriorly and con 
nesting with ejuculatery duct, Eyys, numerous 
in both ascending and descending limbs of 
uterus, messure 110-122 by 73-80 «my; shell 
thin (3-4 ym). 


Relationships: Cardioceplialoides musculosus 
differs from ¢). Aili (ohnston) in the strongly 
muscular nature and bulk of the genital cone, 
und in the presence of “very strong bands of 
longitudinal muscle” in the hindbody (S. J. 


Cardiovephaloides Aili, from Caras novachoilandiae. Second half (measuring 2.1 mm) 


Fig. b- 

of posterior segment, 
Fig. 7, Cardivcephaloides musculoxsusy, from Chlidantas tivhrida. Length 6.3 mm, ventral view, 
Figs, 


8-1), Cardlocephaloldes, evicarpus nsp. Fig. 8---From an unknown host. Holotype: Iength 5.4 


min, ventral view. Fig. 9.-.Skeich of an unmounted specimen, from Phalacrecosnx varius 
Tength 3.46 mim, dorsal view, Fig, 10.—Sketeh of an unmounted spegitnen from an un- 
know! host, Paratype! Length 6.7 mm, luleral view. Fig. 1.—From an unknown host, 
Sagittal section of posterior revion of a paralype, 


ATA OF BIRDS AND MAMMALS 


E 


STRIG 


2% 


=k, 
OFFS, Th 


104 


Johnston 1904, p, 114), Ratio of length of 
Posterior segment/renital cone — 3.0-3.7 in 
Co wiseulosas (averuge 3.3) and 3.45.3 in 
C, Wilt (average 4.8). The two species occur 
in Laylfotines. the first in lems. the second in 
gulls, 


Curdincephaloides avicorpus tsp. 


FIGS, 8-11 
Flous and ovigine Pholacrecarax mielatno- 
leacoy Meevirasiris Gould, frony Dunedin, 
New Zealand, 20.71. /940 (6 specimens, col- 
lected hy Miss M. Fyfe, Otago University). 
P. varins (Gmelin), from Port Gawler, §, 
Aust. 29.91.1998 (several apecimens, From 
two birds), Unknown host (18 attached and 
free specimens) (lype material). 
Aabhirat: intestine. 
Holoiype: length $.4 mm. SAM, E929. 
Paratvpes; E930, (Sections in Neuchatel 
University anid Adelaide University collec- 
tions). 


Description: Body medium-sized, divided into 
distinct anterior and posterior segments. Total 
length 2=7.6 mm. Specimens which have 
recently begun cgg production 2.5-3 mm long, 
Anteriur segment, 0.98-2.40 by 1.40-2.80 mm, 
comprising from three-tenths to four-tenths of 
the body length, subcordiform, with two dilated 
and dorsally incurved lateral expansions in ita 
second half, which appear with their ventral 
connection like a reniform collar surrounding 
base of cephalic cupule; latter spherical ar a 
truncated cone. Posterior segment 1.5-5.4. by 
1.0-2.6 mm, ovoid to spindle-shaped, some- 
times cylindrical (fixed in extended state), 
often slightly arched, dumpy, massive in con- 
tricted state (greatest diam, at Jevel of testes), 
narrower in atviul zone of bursa (0.74~-1.75 
mm) which is delineated by a more or less 
definite constriction at the lust fifth, sixth or 
seventh of the segment, Ratio of hindbady to 
forebody ranging from 1.4—2.5, averaging 1.96. 
Oral sucker (marginal), pharynx and ventral 
sucker all small; ventral sucker a little larger 
than oral, situaled wt mid-length of lorebody 
or in front of it (42-45/100). Oral sucker 
140-200 by 110-200 am, pharynx $5—-180 by 
75-127 am and yential sucker 180-210 hy 
120-200 pm. Cueca enclosed in muscular, 
ventro-lateral bundles and extending ws Fur us 
equator of gentlal cone, i.e. posterior limit of 
vilellaria, Tribocytic organ well developed, 
often protruding anterior io margin of Farebody, 
and peneirating into the lateral expansions. A 
layer oF large cells delimtts the whole active 


GEORGES DUBOIS and L- MADELINE ANGEFI 


surfuce of this organ. Proteolytic glands. dis- 
tributed in nomercus small, relatively seat- 
tered bunches, 4s shown by Bacr (196%, fig. 4}. 
Ovary ovoid. 320 by 230 ym, situated dor- 
sally between 27th and 31st hundredths of 
length of hindholty. Testes multilobed, the mass 
vf which measures 2100 by 1300 jim. Vitel- 
laria ure confined fo hindbouy, profusely de- 
veloped and obscuring its contents; lying in a 
large field along ventro-lateral surface up to 
cunstriclion delineating genital atrium, ¢Last 
follicles 420-890 yim distant from posterior 
end, having their mit between 77th and 85th 
hundredths of this part of body.) Pielel widest 
anterur lo vonnads, where follicles extend 
towards dorsal surface. Vitelline reser yor inter- 
testicular. Mehlis’ cland well developed, Lateral 
and posterior to ovary. Bursa copitlatrix enclos- 
ing a wenital cone, 451-980 by 420-920 jum 
(320-1000 am in diam. inside the atrium). 
Central part of this cone sppears less muscular, 
but more spongy and coloured; its inner wall 
thtown into a few folds delimiting sinuous 
spaces. Uterus extending into first third of 
hindbouy, where it develops several convolu- 
tions, then descends ventrally: behind semingl 
vesicle. it makes a conspicuous right angled 
bend to open inte genital cone. Sinuous ductus 
ejacilatonus, wall of which is 5-16 pm thick. 
opens near entrance of uterus toa genital cone. 
Eggs very numerous, 120-130 by 78-94 am 
(thickness of shell 4—5 pm, up to 8 anv al the 
non-operculate pole), 
Relurionships: The new species resembles 
phyyalis (Lutz, 1926) [syn. ©. safle (Hast- 
Wich, 1954)), as figured hy Duhow (1968, figs. 
187-188) and rediscovered by Buer (1969) in 
the intestinc of a cormorant from Peru 
(Guiahape Islands), but C. physalis differs from 
H In having a much bigger bursa copulatrix, 
eges with yery thick shell and vitellaria extend- 
ing only as far os the beginning of the hursa, 
and in the geographic distribution. 


Cotylurns (Cotylurus) magniacetabuluy psp. 


FIG_ 12 
Host und origin: Cypnur atratus (Latham), 
from Tailem Bend, 5. Aust, 25.%,1945 (16 
specimens). 
iabitar: lower intestine. 
Holotype: Length 2 mm. SAM, E931 (with 
5S paratypes, E932, on same slide; and a 
second slide, E932). 
Descripien, Body very museulur, with 
meridian nyuscles in walls of torebody and 
circular fibres surrounding the oblique open 


STRIGEATA OF BIRDS AND MAMMALS 


ig; at beginning of hindbody are longitudinal 
musckes dorsally, sathered into several bundles 
whieh spread out at level of reproductive 
organs, dorsal one reaching posterior end of 
worm, 

Body 2,0-2.6 mm Jong. Forebody cupuliform, 
hemispherical to spheroidal, obliquely trun- 
cated in front, with yentral border nearly recti- 
lmear and shorter than strongly incurved dor- 
sal border, 0,60-1).71 by 0,740.86 mm, well 
marked off from the gradually attenuated 
cucumiform hindbady, |.34-J.65 by 0.53-0.67 
mm, which is eccentrically fastened to the 
former, Ratio of hindbody to forebody [rom 
2-26, Oral sucker 105-127 by 130-155 ym; 
ventral sucker much larger, cupuliform in pro- 
file, 180-240 by 160-190 um [ratio of the 
average diams, of the latter to the former 1.34— 
1.74, average 1.47); pharynx feebly muscular, 
80-95 hy 64-75 gm. 

Ovary reniform, 130-160 hy 175-210 pm. 
situated between 17th and 24th hundredths of 
hindbody. Testes trilobate, with lobes directed 
posteriorly, the first measuring 210-350 by 
275-320 pm, the second a litle smaller. 210- 
320 by 265-300 pm. Vitelluria very profuse 
through hindbody, and extending anteriorly 
into the two lips of tribocytic organ {erratic 
follicles), Vitelline reservoir and Meblis” gland 
intertesticular. Seminal vesicle lying dorsally; 
ejaculitory duct and uterus (which extends to 
inters¢gmental constriction) uniting and open- 
ing through a short common duct (length 70— 
§0 um) into atrium. Genital bulb, 140-170 
wm in diam., provided with a conspicuows 
muscular thickening at its base, on the dorsal 
side. Eges from 30 to 60 in the uterus, 84-99 
by 58—72 ym {average 89 by 63 pmb. 
Relationships: This new specics closely 
resermbles C. strigenides Dubois, 1958, but 
differs from it in the larger size of the aceta- 
hulum, relative to the oral sucker (ratio of the 
average diams, 1.10-1,13 in C. strigeoidey), in 
the ovary being more distant from the beginning 
of the hindbody (7-8/100 in C. strigesider), 
and in the geographic locality, 


Sehwartzitrema pandubi (Pande, 1939). Dubois, 
1968; 248, figs. 257-260. Dubois & Pear- 


son, 1965; 90, figs. 8, 9, 1967: 190, 
Pande. 1939: 26, figs. 3, 4, 
FIGS. 13, 14 
Hosts and origin: Phalkcrocorax carbo 
(Linn,), from Tailem Bend, S, Aust, 
2biv.l94) (4 specimens), F. gzileirastris 


(Brandty, from Tailem Benu, (54.1941 (16 


205 


specimens, of which LO ovigerous), 154.1941 
wnd =. 26,11.1943) (numerous, ¢ysts, some 
excysting. from stomach, alofg With fish 
remains), 2. relaneleucos (Vieillot), from 
Tallem Bend, 25.x-1945 (14  inmutere 
specimens), 304.1938 and 24,j),/943 (24 
specimens) and €,¥1.1945 (40 specimens), 
flabitat; intestine. 

Descripnan: Body length 0.7-L.4 mm (ovi- 

gerous specimens )- 

Remarks: This parasite appears te be common 

in cormorants at Tailem Bend (cf. Dubois & 

Pearson 1965, 19471. 

The tetrucotyles found in the stomach of P. 
suleiresiris were beth encysted and encapeu- 
lated. The cysts are dark, strong, ovoid, helmet- 
shaped, with a suhconical pole und a circular 
opposite aperture. They measin'e 400-450. by: 
320-360 pam (230-320 jum at the level of the 
opening). The cyst wall varics from 40-90 
gem in thickness. (70-130 ,m at the pole). At 
this stage the structures in the forebody of the 
tetracotyle are clearly differentiated, and the 
conical hindbody ix separated by & murked 
constriction. The characteristic processcs of 
the pscudo-suckers are in & conspicuous posi- 
tion (Fig. 13). The cists are often surrounded 
by « thin layer of hyaline secretion (5-18 ym 
in thickness, 8-24 pm at the pole}. 

The cysts were also found tree in the 
stomach and proventriculus of grebes, Fudire ps 
novaehollandiae Stephens (24.11.1943 and 
14.0v.1943), P. pelioeephalus Jardine & Selby 
(14.1v.1943) and FP. crisvatus = (Linn. 
(24.x1,1947), all from Tailem Bend; from 
the stomach of Pelecanus conspicillatus (Tem- 
minck) along with fish and the crustacean, 
Cherax destructor (2741942, 25.1942, 
17.iv.1944), from Tailem Bend; in the stomach 
of Platcdlea flavipes Gould, slong with fish 
(24.11,1943)}, from Tailem Bend, and from fish 
remains. with escaping moetacercariae. in 
Boraurus poicileptilus (Wagler) (20.v.1949) 
from Mannum, River Murray, S. Aust. 


Family DIPLOSTOMIDAE Poirier 
Subfamily pirLostoMINAE Monticelli 


Baolbophorus confusus (Ktause). Dubois, 1970: 
266, figs, 272-276, 278 (metacercaria). 
Dubois & Peurson, 1965: 95, fig, 13, 

Host and origin: Pelecanus consnicillatus 
(Temminck) from Tailem Bend, 8. Aust, 
April 1938, Dec, 1938, 27.1.1942, 25.-v.1942. 
23 xi.1943, I7iv 1944, Waisesn (40 


2G 


Mature specimens, and 20 metacercariae and 

young specimens). 

Hahita stomach and intestines, 
Deserimion: Body length 1.5-2.2 mm. 
Remarks; The first Australian recurd of this 
cosmopolitan species is that of Diybois & Pear- 
son 41965), who observed un acetubuluni 
measuring 42-65 by 48-92 ,m. In the mature 
specimens of the present collection, this organ 
is smaller, 32-40 by 42-50 p.m, 


2Diplostomam (Diplostomum)  amygdalum 
Dubois & Pearson, 1965: 90, figs. 10-11. 
Dubois, 1970: 300, figs. 297-298. 
Rost vend origin: Nycticorax caledonicus 
(Gmelin), from Tailem Bend, S. Aust, 
1vil940 (9 hinmature specimens). 
Habitat: unknown, 
Dexerinfien; Body length 1.55-0.66 mm. The 
vitcllaria. are not yee developed. This species, 
adapled to the Ardeidae, commonly occtirs in 
Brisbane, Old. ¢ Botanical Gardens). 


Diplostomam (Diplostomum) paryulum n.sp, 
FIGS. 15, 16 
Hosts afd orivin; Hvdraproene  caspia 
(Pallas), from Tailem) Bend, §. Aust., ec. 
1939 (Type material: 5 specimens). Pele- 


ees  conspicillans (Teniminek) — frotn 
Tailem Bend, date? (4 specimens). 
Habitual: intestine. 


Holotype: length 0.56 mm. SAM. E933, 
with 2 of 3 ¢?) purayipes (2934) on same 
slide. 

The type-material was obtained from a Cus- 
pin tern at Tailem Bend, Another collection 
has been found in a pelican from the same 
place: this is probably a case of erratic para- 
sitism. the worms being mixed with metucer- 
caring of Rolhopharus canfusus (Krause). 
Description Body 0.42-0.87 mm long, more 
ar less distinctly divided in two segments. Fore- 
body 0.25-0.53 by 0.24-0.34 mm, oval in out 
line, lolitarm, spoon-shaped with posterior 
margin curved ventrally. Hindhody ovoid, bent 
dorsally, O16-0.94 by 0,21-0.27 mm (dorso- 
ventral diam. 0.21-0.32 nim). Ratio. of length 


GEORGES DUBOIS and L. MADELINE ANGEL 


of second segment to first. from 0.62-0.75 
(average 0.67). Oral sucker rounded, promin- 
ent, 50-62 by 52-63 pm} ventral sucker almost 
equal in size to the oral or smaller, broader 
than long, 45-32 by 55-63 ym, pharynx elon- 
gated, S0-S2 by 24-30 jam (ils antero-posterior 
diath. ds often equal to that of ors) sucker}; 
ogsophagus short, 15-25 um; intestinal caeca 
narrow (5-10 ym), conspicuous in forchody 
and ontering hindbody to terminate not far 
from posterior end. Pscudo-suckers semitunar 
or kidney-shaped, thicker anteriorly, 73-90 by 
by 37-60 pm. Tribocytic organ. approximately 
circular, with a median cleft, 75-100 by 75— 
120 jms proteolytre zland bilobed, with mas- 
sive bean-shaped lobes, lying transversely at 
base of anterior segment. 


Ovary submedian, situated at beginning of 
hindbody, 35 by 60 um. Anterior testit asym- 
metrical, 45-55 by (30-170 um: posterior 
testis bilohed, 60-75 hy 220-250 ,m. Vitellaria 
extending from posterior margin of ventral 
sucker or front of tribocylic organ to caudal 


extremity of body, with the greater density in 


pretesticular zone, reducing beyond to two 
medio-ventral sowings of follicles at level of 
testes. und abutting »gainst the rather compact 
latere-terminal accumulations behind these; 
Vitelline reservoir intertesticular. Bursa copu- 
latrix small, the pore being dorsal and sub- 
terminal (at 50-65 xm from posterior 
extremity); genital cone absent, Eggs few in 
number (1-3), 89-95 by 55-65 pm. 


Relationships: This diplostome is characterized 
by its very small size, being the smallest of 
those described in the subgenus Diplostanuin. 
It closely resembles the South American species 
D, wnucen Szidat, frony Larus dominicanux 
Lichtenstein, but differs ftom it in the size of 
the eggs and io the sclative diameters of the 
oral and ventral suckers, (In the latter, the 
cags measure 110 by 70 pm, and the ventral 
sucker is hirer than the oral.) 


‘there were yet larger diplostomes in the 
collection from HMydroprogne caspia, They 
measure 7 to 1.0 wim in the contracted state. 


Fig, Iz. 
mm. 
Figs. 13, 14, 


Cotyluruy (Cotylirus) magniacctabnles osp., from Cygnus atratus: Holotype: length 2 


Nehwartzitrema pandubi. Fig. 13—Pxcysted terracotyle, from Phalacrocoray meélanvleacas. 


Length 0.47 mm. Fig, 14,—Cyst, from stomach of P.. sulcirestris, 420 by 320 pm- 


Figs 15, (6. 


Diplastamim (Diplostomum) parvulum w.sp. Pig, 15.—From Hydroprogne causpia. Holo- 


type: length 0.56 mm, ventral view, Fie. 16.—From Pelecunay conspicillatus. Length 0.42 


mm., dorsal view, 
Tia, 17, 
Length 1.7 mm, ventral view. 


Viplasramum (Diploviemium) spathageum mbrravense, Tron Larus nevachollandiac. 


STRIGEATA OF BIRDS AND MAMMALS ni 


2S 


Ratio of massive hindbody to forebody from 
0.91-1.25 favernge £02). We would have 
difficulty in describing them. Perhaps they 
helong to the former species, bul the pharynx 
is obviously lacecr (60-70 by 47-52 ,m), 
and the ventral sucker (89 hy 65 xm) ts 
greater than the oral sticker (68-75 by 57-65 
wm), 


Diplostamum  (Diplostomum) — spathaceum 
murrayense (Johnston & Cleland, 1938). 
Johnston & Cleland, 1938: 127, figs, 1-10 
(cercaria). Dubois, 1966a: 40; (970; 341, 
fig. 353, Dubois & Pearson, 1965: 93, fig, 
12. Johnston & Angel. 1941: 140, figs. 1- 
10) flife cycle). Johnsten & Simpson, 
1939: 230, figs. 1-6 (diplostomulum). 


FIG. 17 

Hest unl origit! Larus novachollandiac 

Stephens, from. Swan Reach, River Murray, 

8. Aust, 14.21.1937 (12 specimens); Tailem 

Bend, S. Aust., 3.ii,1940 (8 specimens), 

LQ i.194, (12 specimens}, 19.v.1941 (20 

specimens). 27.1.1942 43 specimens) and 

28.11.1942 (13 specimens); Yalkuri, 8. 

Aust., 24.viii 1957 (one specimen), 

Habitat: intestine. 
Remarks: Johnston & Simpson (1939) believed 
thal the adaly would be found in lariform birds, 
most probably in the silver gull, Larus nevae- 
hwllandiae, However, Johnston & Angel (1941 ) 
reported having found young and adui{t dip- 
lostomes in the marsh tern, Chilidanias hybrida 
(Pallas) [C. leweepareial, aid successfully in- 
fected Limunen fessei Deshayes with eggs 
from the adult flukes. later recovering cer- 
cariae (Cerciria mivrrayensis) from this snail. 
From the occurrences recorded above, it seems 
that the silver gull is an equally, if oot more 
important. definitive host for this parasite. 


Diplostomum (Tylodelphys) podicipinum podi- 
cipinuin Kozicka & Niewiadumska, 1960, 
Dubois, 1970; J8S, figs. 420-421. 


FIG. 18 

Host and origitt Podiceps ertatatus (linn) 

from) “Vailem Bend. S, Aust, 24.xi1947 (16 

(specimens). 

Rabhitat: unknown. 
Description: Body length |.32 im. 
Remarks: ‘Viis is the tirst Australian record 
of this parasite oviginally described from 
Poland, collected from Slovakia and U.S.S.R., 
and characterized by the grea relative 
diamicter of the acetabulum (90 by 95m, 
equal tea quarter of the body breadthi, ¢lon- 


GRORGES DUROIS sad lL. MADELINE ANGEL 


gate pseudo-suckers (170 pm), the rilia of 
length of body to pseudo-aucker (7.7), and 
the presence of a Conspicuous atrin) sphincier, 


Iiysteromorpha platalea Dubinin & Dubinina, 
L940. Dubois, 1970: 397, figs 430-435. 


Diplostomiun ardeiforminun  Odening, 
1962. 
FIG. 19 
Host and erigin: Threskiornis  meliucca 


(Cuvier), from Old., 26.vi.1911 (18 speci- 

mens: Cull. A. Breinl). 

Mabitary unknown. 

In 1840 this minute tremptede was described 
hy Dubinin & Dubinina from the intestine of a 
spoonbill, Platalea leucoredia L,, in USSR. 
{the Volga delta}. Its presence in Indian was 
reported’ by Odening (1962), who considered 
it as a new species designated by the name of 
Diplostomum ardeiformium, from Pseadibis 
papillosa (Temm.). The present record is the 
first Australian reference to this parasite. 
Description; Body length 0.58-0.76 mm, Fore- 
body 0,33-0.40 by 0.36-0.51 mm. Hindbody 
shorily ovoid, 0.23-0.37 by 1.34~0,44 mm, 
scarcely demarcated from forebody by a weak 
constriction. Ratio of hindhody jo forebody 
0,66-0,94. Oral sucker 52-68 by 57-75 ,.m; 
pharynx. 42~52 by 40-45 ym; ventril sucker 
small, almost equal in size to phutyna, 42 47 
by 47-30 um, masked by a well-developed 
tribocytic organ, 210-265 hy 170-225 um, 
Protealyiic gland oval in outline. crescent- 
shaped, strongly lobulated, 95-125 by ILO- 
125 am. 

Ovary 45-55 by 95-135 ym, situated at in- 
terscgmental level. Anterior testis asym 
metrical, 70-105 by 150-210 p,m; posterior 
testis bilobed, 80-125 by 290-390 ym_ Vitel- 
lavia well developed throughout most of 
hody; follicles distributed in the folinceous 
forebouly, expecially at its base, and in triho- 
cytic organ, where they are arranged in the 
form of 4 semicircle: restricted in hindbody vw 
4 medio-ventral ficld, which widens out to 
constitute two Jatero-terminal clusters. Vitel 
linc reservoir intertesticulur. Mehlis’ gland 
lateral, Opposite anterior testis, Eggs, from 3— 
10 in the uterus, 95-112 by 58-68 jon faver- 
age 103 by 62 pm), 

Relationships: Aystercmorpha platatea shows 2 
close resemblance to MW. relloba (Rud.), but the 
Jatter appears to he higger (up to 2.2 mm) 
whereas its eggs are smaller (75-99 hy 48-75 
pm and more numeraus. The acetabulum of 
H_ trilodg 9s always larger than the oral sucker: 
and the hosts are yalious species of cormorants. 


STRIGEATA OF BIRDS AND MAMMALS 


Hysteromerpha triloba 9 (Rudolphi. 1819). 
Dubois, 1970: 400, figs. 436-439. and 
440 (cercuria). T. H, Johnston, 1942: 238, 
Oiplasromu/am cort! Hughes, 1929 (meta- 
cereutis). 

Diplosomum granulosum Goss, 1941. 
Parasirivea slovucica Rysayy, 1958 

Host and origin: Pialacrocoraxk melano- 

fevces (Vieillot), from Tailem Bend, S. 

Aus\,, date? (7 specimens). 

Aabitar: unknown. 

The first record of this cosmopolitan para- 
site is that of Miss O. M, Goss (1941) who 
described it us Diplasioneuen eranulosunt, from 
Pralacrocorax stleiresirix ( Brandt) [= P. ater 
from Perth, W. Aust. T, H. Johnston (1942) 
recorded it from various cormorants, especially 
from Tuilem Bend, 

Deserintion: Body 1,03-1.18 by 0,56-0.67 

mm_ Ventral sucker 90-92 by 94-98 ym: oral 

sucker 70-73 by 83-84 pm; pharynx 50 by 47 

um, Eggs 92-95 hy 63-68 pm. Tribocytic 

organ rounded, funnel-shaped when protruded. 

Proteolytic gland hilobed and trapezoidal. 


Neodiplostomum (Conodiplostomum) brachy- 
orem (Nicoll, 1914), Nicoll, 1914: 346, 
pl. 24, fig. 9. Dubois, 1937; 333, figs. 
11-12; 1970: 418, figs, 451-452. 

Hosts aid origins Ninox novaeseclumdiae 
(Gmelin), from Yalkuri, S. Aust, 
24.vii 1957 (18 specimens, collected by 
W, H, Ewers). Tye alba (Scopoti), from 
Point Turton, Yotke Peninsula, S. Aust. 
12.ix-1970 (3 specimens). 

Habitar: intestine, 

Deseription; Body Jength 1.5-3 min. 

Remarks: This species. is chaructenzed by the 

large size of the ventral sucker, 72-105 by 

73-110 pm (average 87 by 90 .m), and by 

the fact that it occurs im the Strgiformes, The 

testes are larye and symmetrically developed. 

The vitelline follicles have their maximum den- 

sity in the forebody, where they quite often 

reach to the level of the intestinal bifyreation. 


Neodiplostomum (Conodiplostomum) spathula 
australiense Dubois, 1937; 337, figs. 13- 
14: 1970. 428, figs. 465-468. Dubois & 
Pearson, 1967: 96, fig. 7, 
Hosts and origin: Circus approximans Peale. 
from Vailem Bend, S, Aust, Dec. 1938 
{3 specimens), Haliwerus  leucagaster 
(Gmelin), from Wauraltee, Yorke Pen., S, 
Aust, 21-viii 1960 (3 specimens}, Faleo 
peregritins Tunstall, from Naracoorte, S. 


2n9 


Aust, 12.vi.1956 (24 specimens), F. sué- 

niger Gray, from Meningie, 8. Aust., 

&v.1945 (19 specimens), Accipirer cirra- 

cephalus (Vieillot), from Townsville, Qid., 

1911 (one specimen, collected by T. H. 

Johnston). . 

Habitat: duodentim and intestine. 

Descriptions Body length 1.0+-1.75 inm, Vitel- 
laria densest in forebody, sometimes reaching 
the intestinal bifurcation. 
Relationships: The size of the veatwal suckee 
(45-68 by 37-75 pm, average 37 by 67 wm) 
constitutes 4a useful specific character. In this 
respect Neodiplostomum sputhula is distin 
puishable from N. brachyvurum tNicall), 


Neodiptostomum 
latum Tsp. 
FIGS. 20, 21 

Host «and origin: Ninox noyaeseelandiae 

(Gmelin) from Adelaide, S, Aust., April, 

L959 114 specimens), 

Habitat; intestine. 

Holotype: length 1.25 mm. SAM, E935 

with 4 paratypes, E936, and another slide, 

£936, 
Description: Body distinctly bisegmented, 
1.25-1.52 mm long, Forebody flattened. 
lanceolate, 0.79-1.01 by 0.28-0.38 mm, with 
posterior border, where it is wider. curved 
ventrally: Hindbody subcylindrical to claviform, 
always shorter than forebody, 0.40-0.61 by 
0,18-0,21 mm. Ratio of hindbody to forehody 
from 0.50-0,68, averaging 0.56. Oral sucker 
38-47 by 42-50 ;.m; ventral sucker slightly 
larger, 36-52 by 47-57 «om, situated between 
the 50th and Sth htindredths of length of fore- 
bedy, Short prepharynx (10-15 ym); pharynx 
ellipsoidal and musculyr, 37-45 by 26-32 pm; 
ocsophagus reaching length of 40-52 am; cacca 
narrow (about 5-10 ym) im their visible sec 
tion. Tribocytic organ narrowly ellipsoidal or 
almond-shaped. 150-210 by 80-125 pm. 

Ovary oval or rounded, submedian, located at 
beginning of hindbody between 16th and 21st 
hundredths, 55-63 by 63-75 pm. First testis 
appears asymmetrically developed, SU-I1L0 by 
t20-140 pm; second testis clearly bilobed 
{with a postertor median indentation), 80-110 
by 150-185 ,m. Vitelline follicles very con- 
spicuous, with a maximum density at buse of 
forebody; from thence invading wibocytic 
organ and, separaied in longitudinal bands, 
extending beyond ventral sucker, with their 
limit on median fine between the 24th and 
48th hundredths of this part of body; densely 


(Neodiplostomum) laaceo- 


210 


distributed On each side at beginning of hind- 
body. then veeeding from dorsal area 
to hecome 8 wide ventral  cibbon 
which ends in two latero-terminal or sub- 
terminal accumulations, Vilelline reservoir 
yituated at mid-length of hindbody. Mchlis* 
gland lateral, on level with second half of 
first testis, Hermaphrodite canal, which pro- 
longs the incurved uterus, does not traverse a 
genital cone, Genital pore dorsal and sub- 
terminal, at 65-10 pm from posterior ex- 
iremilty of boily. Eggs, few in number (up to 
13}, 94-115 by 63-72 pm (average 104 hy 66 
wm), 

Relationships; Vive species ol the subvenus 
Neodiplosiomum are parasites of night-birds 
and have vitellaria passing beyond the ventral 
sucker; N. americanum Chandler & Rausch, 
N,  cenaiieularain = (Nicoll), N. japerienm 
Dubow, N. rensselati Dubois and N. rravussost 
Duhois, Among them, only the first has an 
ellipsoidal iibocytic organ, but the ovary is 
situaled al the junction of the two segments of 
the body, und the ventral sucker is locutecl 
heiween ont third and two fifths of the Lfore- 
body. This American species reaches 2.9 mm. 


Ncodiplostomum (Neodiplostoumm)  sub- 
aequipartitum Dubois & Pearson, 1967: 
14, fig. 6. Dubois, 1970: 484, fig. 555, 


PIG, 22 
Most and origin: Heliuxter  sphenuriun 
Vieillot), from Tailem Bend. 5S. Aust., Dec. 
1938 (22 specimens}, Is.vil 941 (young 
specimens) 
Hubli: intestine. 


Description: Body 1.16-1.30 mm long, divided 
into two nearly equal segments, Forebody 
0,61-0.68 by 0.36-0.49 mm; hindbody 0.55— 
0.63 by 0.38-0.43 mm, Ratio of the hindbody 
to forcbady from 0.89-1.02 (average 0.93). 
Oral sucker 47-55 by 45 52 «am; pharynx 45-- 
52 by 36-52 pm; ventral sucker a little larger 


GEORGES DUBOIS sod L. MADELINE ANGEL 


than oral, 52-65 by 54-68 pm, situated be: 
tween 45th and 50th hundredths of length of 
forebody; tiboeytc orgun 170-240. by 160- 
210 i. 

Ovary ovoid or ellipsoidal, lying al begine 
ning of hindbody, 95-110 by 130-170 pm, 
Anterior Testis. Igteral, asymmetrical, cunei- 
form or ovoid, [25-150 by 140-210 pm; 
postenor testis bilobed, dumb-bell-shaped, 120— 
150) by 260-320 pm, with greater lobe 
obliquely opposite first testis, Eggs few in num- 
ber (ove to four), 84-92 by 58-65 jum. 


Neodiplostomom (Triloborchidiplostomun) 
diaboli n. subgen., sp. 
FIGS, 23, 24 

Most and origin; — Sarcephilus 

(Boitard). from Tasmania, Oet. 

specimens ), 

Hahiter: unknown, 

Holotype; length 2,05 mm, SAM, F937. 

Paratype in Neuchatel University. 
Description: Body 1,95-2.05 mm long, divided 
by a constriction into anterior and posterior 
segments. Farebody broadly elliptical. spathe- 
shaped, 0.90-0.93 by ©.70-0.73 mm, deeply 
concave posteriorly, with lateral cdges obliquely 
curled ventrad and continuous with each other 
behind tribocytic organ. Hindbady long ovoid. 
or conical, 1.05-1.12 by 0.45-0,56 mim, widest 
at testicular level. Ratio of hindbody to fore- 
budy, 1.2, Oral sucker spheroidal, 90-92 by 
93-95 um; pharynx clliptical in outline, much 
larger than oral sucker and very muscular, 
{22-130 by 118 pm. Ratio of lengths; oral 
sucker + pharynx/forebody, 0,20, Cacca ter- 
Minuting in front of genital atrium, Le. to 
posterior limit of vitellaria, Ventral sucker (ea. 
75 pm) masked by large triboeytic orgun, 460- 
480 by 500-525 ym, irregularly rounded, with 
a very narrow slit; ity frontal border situated 
hetween one sixth and one tenth of the length 
of the forcbody, more or less overlupping the 
pharynx. 


harrisié 
1969 (2 


Fig, 18, 
end with atrial sphincter- 
Fig. 


Diplostornum (Tylodelphys) podticipinum podicipinum, trom Padiceps cristatus. Posterior 


Hivereromorpha platalea, trom Threskierniy melucca, Length 0.64 mm, dorsal yiew. 


Figs, 10,21, Neodiplestomum (Neodiplostomum) lanceofatum n.sp,, from Ninax novaeseelaudine, Fig, 
20.—Holotype: length 1.25 mm, dorsal view, Fig. 21.— Lateral view of hindbody, 
ie. tA. Neudiplostomum (Neodiplasiomum) subacquipartiiin, OQulline of posterior extremity of 


- Neadiplastomum (lrilaberchidiplostamum) diaboti nsp.. trom Sareophilus: hareist), Fig. 


23. Holotype: length 2.05 mm, ventral view. Pig, 24,—Morphology and topography of 


Fin, 22 

hody. 
Tigs. 23, 24 

genital glands of holotype, dorsal view. 
Fin, 14 


min, dorsol view. 


Pharyrgostomuides dasyivi rsp. frum Dagiaris viveryeais (Shaws, Holowge: length W42 


STRIGFATA OF BIRDS AND MAMMALS 21! 


Paes ela ey, 
f 2) 


oS 


4 
Ovary ovoid. transversely clongule, mediah 
ar suhmedian, 130-140 by 180-221) jem, lying 
at junction of forebody and hindbody, Testes 
trilolate (With one dorsal lobe and two Tatero- 
ventral lobes), occupying with ovary, first three 
fifths of segment: anterior testis contiguous 
with ovary, asymmeirical, 140-200 ,tn on left, 
240-320 im on right, and 350-430 um in 
transverse dimension, Second testis 200-230) yam 
and 275-280 yin on two of its lobes, and 330- 
410 »m transversely; posterior border uf this tes- 
tis situated between S6th and 63rd hundredths 
of Iengih of hindbody, Seminal vesicle wel) de- 
veloped, pusitesticular. Vivelline follicles having 
their maximum density at junction of two sey- 
ments and in second half of forehody, pene- 
truting tribocytic organ and extending forward 
to lewe) of posterior border of pharynx; from 
bexinning of hindbody less abundant, covering 
ventral fice in form of two submedian fields 
which widen slightly at level of seminal vesicle 
(distunce from [ast follicles to posterion end 
of the body is 200-270 pm). The anterior 
border of the bursa copulatrix is founct he- 
tween the 87th und the 90th hundredths of 
length of hindbody, The collapsed eggs of the 
paratype measure approx. [25 by 70 pm. 
Relationships: This. parasite [rom the Tas- 
runtan devil is readily distinguished from all 
othe!’ species of the genus Neodiplostomunt hy 
the trilobate shape of the testis®, with the 
exception of Neodiplosromum rantariné Dubois, 
196, of which the pusteriar testty also 
develops three lobes (two Jatera-ventral and 
ene medio-dersal}, This morphological 
character justifies the establishment of a new 
subgenus. For which we propose the name Trilo- 
berchidiplosromum, with the type species N 
(T.) diaboeli nap. A second character com- 
moo to the two species is the ratia of the 
lenpths of oral sucker + pharynx/forebody, 
which varies between 0,20 and 1.25. 
TRILOBORCHIDIPLOSTOMUM 4. subgen. 
Devnosgds: Neadiplastomum with the iwo 
fesles ur only the second testis trilobate. Com- 
plex of oral sucker and pharynx usually he- 
tween one-lifihh and one-quarter of length of 
forebody. Tribocytic organ tending to hyper- 
trophy. Intestinal parasiles of mammals. 


GEORGES DUBOIS and L. MADELINE ANGEL 


Type spéecles: N. (T.) didbofi nsp., in Sar- 
cophiluy harrixii (Boitard). ftom Tasmania. 
Consubpeneric species: Ny CT.) ramoriat 
Dubois, 1966 in Leantocebus  nigricatliy 
(Spix), from South America, 


Neodiplostomulum sp. 

floxty and origin: Netechiy scutuney (Peters) 
{Cphidial, from Tailem Bend, S. Aust,, 
I.v,1940 (8 specimens and 2 cysts), Pyea- 
decihis porphyriacusy (Shaw) JOphidial, 
from Adelatde Zoo, S.xt.i957 (5 speci 
mens). 

Rahitat; digestive traci, anu subperiioneum. 


Deseriplien: Body ova). 255-475 by 210-285 
pm. Cacca gradually and irregularly distended, 
filled with a yellowish substaiice, Cysts oval, 
350-380 by 300-320 pm, Cyst wall fibrous, 
15-40. ym thick, 


Remurks: One specimen only of this meta- 
cervaria (335 by 245 am) was fourd in the 
intestine of Grallina cyanoleaca (Latham) 
(Passeriformes) [rom Tailem Bend, 22,1,1943, 


Posthodiplostomum ausirole Dubois, 19357: 
1970: 510, figs. 585-588, Duhois & Pear- 
som. TY67! 201, fie ¥. 

Rosts and origins Phalacrocerax sulcirestris 
(Brandt), from Vailem Gend, $. Aust. 
26.11.1943 (2 specimens), P. saelanoleucos 
(Viellor}, From Tailem Bend, 6.vi.1945, 
31.1948, Oct. 1960 (27 specimens) and 
25.x.1945 (4 young specimens). Pelecanuy 
couspletlaius (Temminck), from Tailem 
Hen, date? (7 specimens), Aydroprogie 
casela (Pallas) from Tailem Bend, Dee. 
1939 (2 specimens}. Kgrevia alba (Linn.), 
from Tallem Bend, June 1937 (one speci- 
men). 4rdea novaehollandiae Latham, trom 
Tailem Bend, |v.1940. 24.41.1943, 6-vi.1945 
and 31.V.1949 (19 Specimens); Swan Reach, 
5. Aust., 15.xi.1937 (14 specimens); Tailem 
Bend, May 1938, 1.v.1940), 24.11.1943. (27 
young specimens cseaping and escaped from 
cvsts}. Nycticarex caledomicus (CGrnelin) 
from Mary River, Northern Territory, May 
1962 (17 specimens), 

Habitat: Stomach and intestine. 


2 AS in severul species of the subgenus Peralaria Keatise, In particular Aferia (Paralaria) pseido- 
chathratd Keause, A. (P.} mustelae mustelae Bustna. and especially 4. (P.) mustelae canadensiy Web- 
ster anu’ Wolfgang (cf. Dubois 1970. figs. 699-700), 

In Frhvinela snareophile Sandars 1957, the posterior testis is “characterislically bilubed”. The fure 
body is “typically longer than the hindbody, and the tribocyltic organ “is usually between one- 
quarter and one-third of Ure Jenpth of the anterior scxment’” 


STKIGEATA OF BIRDS AND MAMMALS Fa 


Deseription: This common species is charac- 
terized by its small size: the ovigerous speci- 
mens measure (1.42-98 mm [fine to nine 
eggs ih uterus). Ti life, there is “orange colour 
around miusucker”. Gral sucker 28-31 hy 30- 
37 jum: pharynx 26-30 by 19-25 ,«m. vencral 
sucker 42-68 by 45-65 ym. 

Ovary situated at beginning of posterior 
segment. First testis asymmetrically developed; 
second testis bilobed, sub-cordiform, with: an 
anterior concavity, Vitellaria very dense at 
base of forebody, extending anteriorly beyond 
ventral sucker, in some specimens to a level 
about equidistant from ventral and oral 
suckers: in hindbody, extending on ventral side 
to pasterior border of second testis, or only to 
intertesticuliur region, Uterus. ventral, bending 
dersad uotil it arrives anterior to the bursa 
copulatrix, and then enters genital cone, Eggs 
73-94 by 48-63 wnt (average 82 by 54 um), 

Cyst transparent, ellipsoidal, 340 by 220 
pom, having a thin wall (6 to 8B ym in thick- 
ness}, 


Subfamily anammnan Hall & Wigdor 


Fibricola intermedius (Pearson, (959), Pearson, 
1959; 311, figs. 1-8. Dubois, 1970: 437, 
fig. 72%, 
Haw anid orfgink Bydromys chrysagaxter 
Geoffroy, from River Torrens, Adélatde, S. 
Aust, July 1923 (21 specimens, some of 
which have one exe in the uterus). 
Hahitar intestine. 
Description; Boy length 0,75-0.98 mam. 
Relationships: This species, the lype-host of 
which is Ratins assimilis, was found in the 
water rat, Hydroniys chrysogaster. by Pearson 
(1961). [t differs from F. minor Dubois (also 
from HA. cliryvoguster) in having erratic vitel- 
line follicles in the hindbody: they form two 
distinct lateral bands which extend as far as 
the zone of the second testis, and even beyond. 


Pharyngostomoides dasyuri nsp- 
FIG. 23 
Hoss and = arigin: Dasvurus  Viverriine 
(Shaw) from Teena Estate. Tas, 9,47.1966 
(3 specimens). 
FAlabitat: small intestine, 
FHfolotype: length 0.62 mm, SAM, E938, 
paratypes ¢3 slides} E929, 
Description; Body oval, tnilistinetly biseg- 
mented. O.58-6.67 mm long. Forebody mar- 
supiform, 0.33-0.40 by 0,45-0.48 mim (when 
contracted), with Jateral and posterior margins 


folded ventrally. Hindbody more or lesa coni- 
eal, 0.23-0.27 by 0.35-0.41 mm (when con- 
tracted), Oral sucker subterminal, 65-738 by 
75-94 wm; pharynx ellipsoidal, more muscular 
buat smaller than oral sucker. 57-63 by 40-48 
wm ventral sucker bigeer, elliptical tn outline, 
SR-78 hy 95-110 jum, usually partially covered 
by uibocytic organ: oesophagus shorl, Pseudo- 
suckers cupuliform, 68-75 by 60-63 ym. Tri- 
bocytic organ well deycloped, elliptical in out- 
line, 155-195 by 125-150 am, with longitud- 
inal slit-like opening, 

Ovary ovoid. 30-75 by 85-LID ym, sub- 

median, located dorsally at base of forcbody. 
Testes rounded of ovold, approximately equal 
in size, 100-140 by 130-180 ym, situated side 
by side in Anterior part of hindbody, close 
together. Seminal vesicle well developed. post- 
testicular. Vitelline follicles confined to fare- 
body, with a dense distribution from Jevel of 
anterior murgins of testes, gradually decreas- 
ing forwards, penetrating tribocytic organ and 
extending medially almost to bifurcation of 
oesophagus, laterally to level of ventral sucker. 
Vitelline reservoir pretesticulur, median or 
suhbmedian, at junction of anterior and pos- 
terior segments, Eggs 115-118 hy 60-6] pm. 
Fresh material is needed for a description of 
the bursa copulatrix. 
Addendum, On 23rd June, 19723, Dr, G. 
Gregory collected 64 uncontracted specimens 
from the intestine of Daryurur miaculutus 
(Kerr), at Ben Nevis, Tusmania, These are 
described below, 

Body length 1,22~1.53 mn. Forebody coch- 
leariform 0.68-0.90 by 0,73-0.95 mm, Hind- 
body conical, rounded at extremity, (),.52-0:71 
by 0.50-0.71 mm at level of testes, Ventral 
sucker 75-104 by 95-117 um (average 90 by 
130 gamj, subequal to or larger than oral 
sucker, AS—95 by 8S—J15 pm (average 80 by 
95 zm}, but more muscular, Pseudo-suckers 
cupuliform, with prosdetic glands well devel- 
oped. Tribocytic organ 300-340 by 210-300 
am (230-350 hy 265-430 when com- 
pletely protruded). Pharynx small, 55-73 hy 
52-65 «wm. 

Ovary submedian, 110-135 by 120-160 um, 
Testes spherical or ovoid, situated side by side, 
180-255 by 200-300 pm, Seminal vesicle 
inter- and post-testicular, discharging through 
muscular ejaculatory duct (ejaculatory pouch 
absent), which unites. with descending limb of 
uterus, (Ascending limb of latter reaching 
mid-portion of tribacytic organ.) Vitelline fol- 
licles confined to forgbody, distributed in two 


214 


lsleral (asses confluent anteriorly at ace- 
tabular level, divergent backwards where they 
terminate in contact with the testes. extending 
medially to level of intestinal bifyrcation (jist 
posterior to pharynx). Tn the triangular space 
between these two masses are the Jonyitudinal 
aperture of tribocytic organ, the ovary and 
the vitelline reservoir (median and pretesticu- 
lar). Bursa copulatrix muscular, rounded, 
220-300 pm in diam., occupying second half 
of posterior segment, with dorsal, subterminal 
aperture and deep genital atrium, Eggs | to 
15, 110-130 by 65-78 pm (largest 130 by 78 
pm), average L118 by 71 um, 

Relationships;  Pharyrgestomoides — daxyuri 
nap. seems to be closely related to P. procyonis 
Harkema, 1942, but dilfers in the smaller body, 
the inequality of the suckers, the position of 
the ovary, the absence of ejaculatory pouch, 
the extension of the vitellania up to level of 
intestinal bifurcation, the greater dimensiuns 
of the epgs, and the geographic distn- 
bution, It is probably identical with Pharyn- 
roxtomoides sp, of Sandars (1957. p, 263), 
recoyered from the intestine of a Tasmanian 
tiger cat, Dasyurus maculatus Kerr). 


Kanily PROHEMISTOMIDAE (Dubois, 1938) 
Sudarikoy, 1941 


Subfamily PROHEMISTOMINAF Lutz 


Mesusteplianus haliasturist Tubsangui & Masi- 
tungan, 1941; 138, pl. 3_ fig. 3. Dubois & 
Tearson, 1965: 96, fig 14 (from Aaliay- 
tur); 1967; 202 (from Pelecanus). 
Mesosirphanus minor Dubois & Pearson, 
1965. 

Host and origin: Pelecenus conspivillatas 
{Temminck), from Tailem Bend, S. Aust, 
dale? (24 macerated specimens), 

Hubitai. unknown. 


Toybangut & Masilufigan recorded this species 


GEORGES DUROTS and 1. MADELINE ANGEL 


from: the small intestine of Hauliestur internivedins 
Blyth from the Pampanga Province (Luzon: 
Philippines). Dubois & Peurson (1963) re- 
described it by the name of Mesostephenus 
miner from Aaliastur sphenurus (Vicillov) 
from Brisbune, Qld., and subsequently (1967) 
from Pefecaunus conspicillaras (Temminck) and 
Anhinga novachollandiae (Gould), from 
Mackay and Kola, Qld. The fishing-kite is 
probably un occasional host, 
Deseription: Body oval, with small caudal ap- 
pendix, 0.9-1.5 mm long by 0.3-0.5 mm in 
maximum width; anterior part well expanded, 
slightly concave ventrally, with lateral borders 
more or Jess rolicd up into a gutter and mect- 
ing posteriorly. Oral sucker 38-52 »m,; ventral 
sucker dightly larger, 37-35 by 40-60 ,»m, 
situated between che 40th and 45th hundredths 
of the length of the body; pharryns. ellipsoidal. 
32-50 by 24-40 pm; oesophagus 47-52 am 
long, Tribocytic organ oval in shape, 200-210 
by 150-160 «jm, with a longitudinal slit. 
Ovury globular, 60-80 ym, level with second 
half of anterior testis, slightly to one side of 
median Jine, opposite cies sac. Testes sub- 
lobular to ovoid, close behind one another, 
111-190 by 90-150 ym. Vitellaria compused 
of fairly large follicles disposed in at ecoen- 
inc wreath (diameter 300-520 jm) around 
tribocytic organ. The two characteristics of the 
species are chat vitellaria (1) do not reach 
acetabular level (limit 44th to 50th hundredths 
of length of body, ie, distant 28-35 ym from 
posterior border of ventral sucker), and (2) 
overlap only first balf of posterior testis. Ratio 
of the length of the body to the diam. of the 
vitelline wreath ranging from 2,7-3.2 (average 
3). Cirrus sac well developed, elongated, club- 
shaped, 310-530 by 50 100 ym, extending 
fornvards to zone of first testis or even. beyond. 
Uterus short, with vaginal sphincter conspic- 
uous. 20-40 by 29-55 ym. Egps one to six in 
number, 90 99 by 65.73 um. 


References 


Base. J. G, (1964)—Un ‘Trématode parasite du 
Cormoran Phedlacrocorax bougainvilleti 
(Less.) cles fles Guanape, Pérou, Pargzit, Sb, 
24, 7 15. 


Cease. E, E (1971).—A new avian trematode. 
Prov. Linn, Suc, NSW. a5, 500-504. 


DUKENIN, V. B., and DWHININA, MON. (1840). — 
iParasitofatinc Ucs Otseaux coloniaux des 


réserves de Is région d'Astrakan|. 
axtrakh. ges. Zakhov 3, 19O-29R. . 

Dubos, G, 41937), —Etude de quelques Strides 
WAustralie ef nate sur le genre FrAvicata 
Dubois 1922, Annis, Parasir. hum. camy,, 15, 
(3 4). 231-247, 333-353. 

Dinos, G. (1938)—Monographic des Striaeida 
(Trematoda), Mém-. Sve. aenchdt. Sci. nat. & 
535 pp. 


Trudy, 


* Incorrect original spelling Swliasfurns. 


STRIGEATA OF BIRDS AND MAMMALS 


Dunas, G. (1966a).—Du_ statut de quelques 
Strigeata La Rue, 1926 (Trematoda). II. 
[German and English summacies], Bull, Sec, 
neuchdtel, Sci, nat, 89. 19-56. 

Dupors. G. (1966b),—Un Néodiplostome (Tre- 
matoda; Diplostomatidae) chez je tamarin 
Leontocebus nigricollis (Spix), Revue suisse 
Zool, 73 (2), 37-42. 

Durois, G, (1968).—Synopsis des Strigeidae et. 
des Diplostomatidae (Trematoda). Premiére 
Partie, Mém. Soc. neuchdt, Sci. nat. 10 (1), 
1-258. 

Durots, G. (1970).—Synopsis des Strigeidae et 
des Diplostomatidae (Trematoda). Deuxiéme 
Partie, Mém. Soc.. neuchdt. Sci. nat. 10 (2), 
259-727, 

Duzors, G., & Pearson, J. C, (1965).—Quelques 
Strigcida (Trematoda) d’Australie. Brit. Soc. 
neuchatel. Sci. nat. 88, 77-99. 

Durots, G., & Pearson, J, C. (1967)—Quelques 
Sirigeida (Trematoda) d’Australie. II, Tbid., 
90, 185-204. 


Goss, O. M. (1941).— -Platyhelminth and Acan- 
thovephalam parasites of Jocal shags. J. R. 
Suc, W, Aust. 26, 1-14. 

HartwicH, G. (1954).—Darmhelminthen von 
Larus modestus und Puffinuy griseus aus 
Peru, Betr. Vogelk., 3 (6), 258-270. 


Jounstos, S. J. (1904).—Contributions ta a 
knowledge of Australian entozoa, No, 1IT.— 
On some species of Holostomidae from Aus- 
ayes birds. Proc. Linn, Sec. NSW, 29, 
108-116. 


JouNston, T. H, (1942).—Trematodes from Aus- 
tralian birds. J. Cormorants and darters. 
Trans. R, Sac. 8. Aust: 66, 226-242, 


Jounston, T, H. & Anaer. L. M. (1941)—Life 
cycle of the trematade, Diplostomim murray- 


215 


ense J. & C, Trans. R. Sac. S. Aust. 65, 
140-144. 

Jounston, T. H., & ANGEL, L. M, (1951),—The 
morphology and life cycle of the trematode. 
Apatemon tntermedius, from the black swan. 
Trans, R. Soc. S. Aust. 74, 66-78. 

Jounston, T. H., & CLELAND, EF. R. (1938)— 
Larval trematodes from Australian terres- 
trial and freshwater molluscs. Part TV. Cer- 
caria (Furcecercaria) murrayensis n.sp. Trans, 
R. Soc. S. Aust, 62 (1), 127-131. 

Jonnsron, T. H., & Simpson, E. R_ (1939).—The 
diplostomulum stage of Cercaria murravensiy. 
Trans, R, Soc, §. Aust, 63, 230-237. 

Nico, W. (1914).—The trematode parasites of 
North Queensland I. Parasitology 6, 333-350. 

OnENING, K. (1962).—Trematoden aus Indischen 
Vogela des Berliner Tierparks, 4. Purastikde. 
21, 381-425, 

Panve, B. P. (1939).—Two new species of tre- 
matodes from Anhinga melanogaster, the 
Indian darter ot snake-bird. Proe. natn. Acad. 
Sci, India 9, 22-28. 

Pearson, J, C. (1959),—Neodiplastomum inter- 
medium nsp. fram the allied rat, Rastus 
assimilis, with remarks on the genera Neadip- 
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Pearson, J. C. (1961)—Observations. on the mor- 
phology and life cycle of Neodiplostomum 
intermedium (Trematoda: Diplastomatidae). 
Ibid., 51, 133-172. 

Sanpars, D, F, (1957).—A new strigeid trematode 
from an Australian marsupial. J, Helminth. 
31, 257-264, 

Tupanaut, M. A,, and MAsiLunaaw, V, A. (1941). 
—-Trematode parasites of Philippine verte- 
brates, IX. Flukes from the domestic fowl 
and other birds. Philipp. J. Sci. 75, 131-140, 


OBITURARY: KEITH RODNEY MILES 


Summary 


Keith Rodney Miles died on 25" March 1972 after an illness which took him intermittently from 
duty over the previous six months. He was born in Western Australia on 8" April, 1915. 


EE 


KEITH RODNEY MILES, 


D.SC., F.G.S. 


OBITUARY 
KEITH RODNEY MILES, D.Sc. F.GS. 


Keith Rodney Miles died on 25th March, 
1972 after an illness which took him inter- 
mittently from duty aver the previous six 
months. He was bora in Western Australia on 
8th April, 1915. 


He completed his formal education ia Perth. 
graduating B.Sc. in 1937 from the University 
of Western Australia, with Honours in Econo- 
mic Geology. He was also at this time lecturing 
in the School of Geology, 


The neXt six years was spent on the staff of 
the Geological Survey of Western Australia, 
during which time he completed a thesis for 
the degree of Doctor of Science, to which he 
was admitted in 1942, His thesis covered the 
jasper bars and banded iron formations: of 
Western Australia. 


Miles joined the Geological Survey of South 
Australia in 1944, This was the beginning of 
the period of growth and development of the 
Survey under §, B. Dickinson, and Miles was 
the first of a group of new appointees compris- 
ing in addition T. A. Barnes, R, C. Sprigg, and 
the present writer. 


He spent some ten years with the South 
Australian Survey at this stage and left in 1954, 
firstly to accept an appointment as Deputy 
Chief Geologist with Australasian Oil EXplora- 
Uon Limited and suhsequently for private prac- 
tice as 4 consultant. He rejoined the Depart- 
ment of Mines in 1963 as Chief Geologist, 
hecame Deputy Director in 1970 and al the 
time of his death was Acting Director of 
Mines. 


Miles’ professional career took him through 
a very wide range of geological interesis, His 
first project—the ironstones—led him Jater to 
a detailed study of the Middleback Ranges, fo}- 
lowed by a Bulletin on this subject. We under- 
took many hard-rock mineral exploration pro- 
jects. several maior dar site studies, and geo- 
hydrology investigations, He travelled widely 
throughout Australia and New Zealand and 
undertook an extensive overseas assignment 
for the Government in 1970, 


Miles was responsible for some sixty pub- 
lished works, including three major Bulletins of 
the Geological Survey of South Australia, and 
a great many unpublished reports A selected 
bibliography is given below. 


Miles Was a conscicntious member of many 
professional organisations and made « contri- 
bution at Committee and executive level when- 
ever called upon to do so, He waa a Fellow of 
the Geological Society of Londen, a Member 
of the Geological Society of America, 4 Mem- 
ber of the Society of Economic Geologists, a 
Member of The Australasian Institute of Min- 
ing and Metallurgy and a past Chairman of the 
Adelaide Branch, a member of the Geotogical 
Society of Australia, a member of the Profes- 
sional Division of the Australian Petroleum 
Exploration Association, He was a Fellow (Life 
Member) of the Royal Society of South Aus- 
tralia and served as a Council member from 
1963-66, Vice President 1966-67 and 1968-69, 
and President 1967-68. 


So much is the offictal record, 


As a colleague, close friend and confidant 
of nearly thirty years, Keith Miles’ death is a 
personal loss, Much more thin this however, is 
the Joss of his experience and judgement at 
the moment when he was in a position to offer 
these to greatest effects at both administrative 
and scientific levels. Keith was fiercely loyul to 
his colleagues and staff und took preat care to 
ensure that those for whom he Was respon- 
sible had every opportunity for scientific de- 
velopment. 


He was jeatous for the status of the profes- 
sion of Geology and set the highest standards 
at probity and workmanship in all that he 
undertook and in his dealings with his fellows. 


A geological family becomes accustomed 
over the years to frequent and sometimes long 
periods of separation, We offer to his wife and 
daughter sincerest sympathy for this the Jang- 
est and most difficult separation of all. 


L. W. Parkin. 


eee 
Trans, R. Soc, §, Aust. Vol. 96, Part 4, 30 November 1972. 


UK 


1937 


1y38 


1938 


1938 


1938 


1938 


1939 


1940) 


1940 


4 


1942 


1942 


1942 


1942 


1942 


1942 


1942 


1942 


1943 
1943 


Bibliography 


Lancefield Group, Report on Betia main 
lode. GUM.L. 22161, Mt, Margarct G.F 
Aun, Rep. geol, Surv. West Aasi. 1938, 
2-71 


The geology and physiography of the 
Lower Chitlering area, J, Prac, RB. Sec, 
Wesr, “lust. 24, 13-41. 

Gladiator {lace Augusia} Gold Mines, Mt. 
Margaret goldfield. Ann. Rep geol. Sir. 
West, dist. 1938, 27-28. 

The Mury Mac Gold Mines, G.M.L. 2206T, 
Laverton, dan, Rep. geal. Surv. West, 
Aust, VO38, 28-29, 

Notes on the geological struciure of por- 
tion of the Mt, Margaret Goldficld. .4nn. 
Rep. geol, Surv. West, Aust. 1938, 29-41. 
Notes on banded Jaspillites of the ML. Mor- 
gans—Mt. Murgatet District. Mt, Margaret 
Goldfield. Ann. Rep. geal, Surv. West, Aust, 
1938, 31-32. 


Notes on some mining groups in the Mt. 
Margaret G.Fo dian, Rep. geal, Surv. West. 
Aust. 1939, 43-56. 

Notes on “Platy” and “Massive” types of 
Jaspillites from Evanston. Avi. Rep, geol. 
Surv. West, Aust, 1940, 17-18. 

Sumomiary of petrological notes on same 
rocks from the Yilgarn Goldfield. Ant. Rep. 
geal. Surv. West, Aust, 1940, 18. 


Magnetite-hemiatile relations in the banded 


iron formations of Western Australia. 
Prac. Australay, fast. Min, Merall, 124, 
193-201. 


Repurt om a petrological Investiganion of 
nictasomatism neur (fie Corinthian ore body, 
Aan. Rep. peal, Surv. West, Aust, 1942, 7. 


The geology of Tindals. Coofgardie Gold- 
field. Anni. Rep. feel, Saev, West. Aust. 
1942, 7, 


Two reputed {ron are deposits in the vicinity 
of Albany, S.W. 4nn. Rep. geol, Siev, West. 
Aus 1947, 7 

The Hill 60 lode, Mi, Magnet Gold Mines 
Lid. Mt. Magnel. 4nn. Rep. geal. Surv. 
West. Aust, 1942, 8 

A supposed manganese and hematite de- 
posit near Wallangie, Coolgardie Goldfield. 
Ann, Rep. weol. Sury, West, Atet, 1942, 8, 
Geological notes on boring in the Mt. 
Palmcr district. Yilgarn GF, dyn, Rep. 
geol. Surv. Went. Aust. 1941, 8 
Mctasomatism near the Corinthian ore 
budy, Western Australia, Prac. Austrafay, 
Inst. Min, Metall. 125, 71-85. 

The blue asbestos bearing banded iran forr- 
mations of the Hamersicy Ranges, W.A. 
Biull, geal, Surv. West, Aust. 100, 4-37. 


Gruncrite in Western Australia, ert. Miner. 


4 2°08. 


Jasper bars and ccopomic geology in Wes- 
tern Australia. Frac. Atestralas. [nxt. Min. 
Metali. 131-132, 187-2416, 


1943 


1943 


1943 


1943 


1943 


19.43 


1944 


1944 


1944 


1945 


1945 


1945 


1945. 


1946 


1946 


1446 


1947 


19.47 


1948 


1949 


Some kyanite-bearing tocks From. the East- 
ern Goldficlds, Western Australia. J. Pree. 
R, Sov, West. Aust, 27, Y-25, 

Bentonite at Mardragee. Western Australiu 
Ann. Rep. geol, Surv, West, Awar, 1943, 24, 
Notes on a salt. depovit near Wyalkutebem. 
S.W. Div. Aue, Rep, geol, Surv, West, Aust. 
1943, 24 


Notes on the Salt Lakes—Compion District. 


Ann. Rep. geol. Surv. West, Aust, (43, 
24-25, 
Sctheelite deposits in the Central Goldfields, 


Western Australia, Ann, Rep. geel. Surv. 
West, ust, 1943. 25-29, 

A lepidolite depasit near Tantalire Hill, 
Londonderry, Coolgurdie G.F. Ann. Rep. 
geol, Surv. West, Aust, 1943, 29-30. 
Notes on un inspection of quiriz crystal 
deposits, Goomalling, Moraws and Payne's 
Find. dra. Rep. geal, Surv. West. Anse, 
1944, 50, 

Preliminary report on limestone prospects 
—Commonwealth lund south of Fremanitle. 
Ann. Rep. peal, Surv, West, Ast. 1944. 
50-53. 


Notes on sampling Douro Rd. Limestone 
quurry, South Freemunile. Ann. Rep. eevl. 
Surv. West, Aust, 1944, 53, 


Report on the Hill 60 lode. Mi, Magnet 
Gold Mines Ltd. Mt. Magnet. dain. Rep. 
weof. Surv. West, Ast, 1945, 41-43. 
Geologicul notes on boring in the Ml. 
Palmer district, Yilegarn GF, ann. Rep. 
geal. Surv. West, Aast. 1945, 43-46. 
Report an the geology of Tindals, Cool- 
gardie G.F. Ann. Rep. geol, Sire, Wes, 
Aust, 1945, 47.56. 

‘Vantalum and niobium, Rall, Dep, Mines 
Miner. Resour, West. Aust. 3, (with D. 
Carroll & H. P. Rowledge), 


Metasomalism of jasper bars of Western 
Ausiralia, Quart. #. geal Soc. Lend. (02, 
115-155, 

Investigation of some phosphatic nodules 
from Wandaragan, Western Australia. 
Proc. R. Suc, West, Awal. W, 75-82. 
Limestone deposits at Hallett Cove. Min. 
Rev. Adelaide, 82. 68-75. 

Precambrian granites and grumitixation with 
special reference to Western Australia and 
South Australia. Trey. Ro See. 9. Aust. 71, 
54-66, 

fa: Matheson, R. 8. The Mining Groups of 
the Yilgarn Goldfield, north of the Great 
Eastern Railway. Ha/l. geal, Surv. Weer. 
Ausi., 101, 9, 15, 27, 29-31, 41, 54. 216- 
342. 

In: Matheson, R. 8. The Dandaragan phos- 
phate deposits. Gall. Dep. Mines. Miner. 
Resour, West, Aust, 4, 

fn: Hobson, R. W. & Muthesan, R. S&.. 
Greenbushes mineral field, Bell, geo?, Surv, 
Wot test, 102, Th. 42-58, Bd-85_ 123, 
195, 202, 27. 


1949 


1950 


1950 


1950 


1950 


1950 


1951 


1952 


1952 


1952 


1952 


OBITUARY—KEITH RODNEY MILES 


Diamond drilling marble deposits at Penrice 
near Angaston, Min. Rev. Adelaide, 88, 
103-134, 

Use of banded iron formations in a geolo- 
gical approach to prospecting in the Western 
Australian Goldfields. Proc. Australas. Inst, 
Min. Metall. 156-157, 47-53. 

The geology of the South Para Dam pro- 
ject. Bull. geol. Surv, 8, Aust. 24, 

In; Johnston, W., A geological reconnais- 
sance survey. Bull. geol. Surv. West. Aust. 
106, 55-56. 


Geology of portion of the Mt. Margaret 
G.F., and, Garnetised gabbors from the 
Eulaminna district, Mt. Margaret G.F. 
Bull. geol. Surv. West, Aust. 103° (with 
R. W. Hobson & R, S. Matheson). 


Moulding sands. Aull. Dep. Mines Miner. 
Resour. West. Aust. 5 (with H. A. 
Stephens), 


Origin and salinity distribution of artesian 
waler in the Adelaide Plains, South Aus- 
tralia, Econ. Geol. 46, 193-207, 


Tertiary faulting in northeastern Eyre Pen- 
insula, South Australia. Trans. R. Soc. S. 
Aust. 75, 89-96. 

Geology and underground water resources 
of the Adelaide Plains area, Bull. geol. 
Surv. S. Aust. 27. 

Middleback map sheet, Geological atlas of 
South Australia, 1:63,360 series. (Gcol. 
Surv. S. Aust.: Adelaide.) 

McGregor map sheet, Geological atlas of 
South Australia, 1:63,360) series. (Geol. 


1952 


1953 


1953 


1953 


1953 


1954 


1954 


1969 


219 


Surv. S. Aust.; Adelaide.) (With J. R. 
Eley, ) 

Roopena map sheet, Geological atlas of 
South Australia. 1:63,360 series. (Geol. 
Sury, §. Aust.: Adelaide.) (With R. K. 


Johns & M. Solomon), 


Banded iron formations in Western Aus- 
tralia. Proc, Fifth Emp. Min. Metall, Cang, 
Aust, N.Z, 1, 59-71. 


Koolyanobbing iron ore. Proc. Fifth Emp. 
Min. Metall, Cong. Aust. N.Z. 1, (72-176. 
Wilgic Mia Weld Range iron ore. Proc. 
Fifth Emp. Min. Metall. Cong. Aust. N-Z. 
1. 242-244. 


Iron ores of the Middleback Ranges, South 
Australia. Im Geology of Australian Ore 
Deposits. Proc. Fifth Emp. Min. Metall. 
Cong, Aust. N.Z. 1, 449-463. (With E. A. 
Rudd.) 


In McGrath, J. C. & Gray. N. M., The 
geology of the country about Coolgardie, 
Coolgardie. G.F., W.A. Bull. geol. Surv. 
West. Aust. 107, 58, 78. 


The geology and iron ore resnutces of the 
Middleback Range area. Bull. geol. Surv. S. 
Aust. 33, 


Geological conditions and arbitration of 
dredged channel and turning basin, Bluff, 
New Zealand. Jn G, A. Kiersch & A. B. 
Cleaves (Eds.), “Engineering Geology Case 
History No, 7 Legal Aspects of Geology in 
Engineering Practice”, pp- 21-32. (Division 
on Engineering Geology of the Geological 
Society of America: Colorado.) 

H. WOPFNER. 


ROYAL SOCIETY OF SOUTH AUSTRALIA INCORPORATED 


Patron: 


HIS EXCELLENCY THE GOVERNOR OF SOUTH AUSTRALIA, 
SIR MARK OLIPHANT, K.B.E. 


OFFICERS FOR 1972-73 


President: 
K. E. LEE, D.Sc. 


Vice-Presidents: 


H. WOPFNER, Ph.D. G. F. GROSS, M.Sc. 3 
Secretary: Treasurer: 
C. J. M. GLOVER, M.Sc. S. A. SHEPHERD, B.A., LL.B. 
Editor: Assistant Editor: 
H. B. S. WOMERSLEY, Ph.D., D.Sc. W. K. HARRIS, M.Sc. | 
Librarian: Programme Secretary: . 
N. H. LUDBROOK, M.A., Ph.D., J. CARRICK, B.Sc. 
D.LC., F.GS. 


Members of Council: 


Hj. EICHLER, Dr.rer.nat. J. W. HOLMES, M.Sc. sf 
P. J. M. GREENSLADE, M.A., Ph.D., I. M. THOMAS, M.Sc., M.I.Biol. 
D.LC. M. J. TYLER 


R. H. FISHER, A.U.A. 


Auditors: 
Messrs. MILNE, STEVENS, SEARCY & CO. 


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