VOL. 97, PART 1 28 FEBRUARY, 1973
TRANSACTIONS OF THE
ROYAL SOCIETY
OF SOUTH AUSTRALIA
INCORPORATED
CONTENTS
Alley, N. F. Landsurface Development in the Mid North of South Australia 1
Edmonds, S. J. Australian Acanthocephala, No. 14. On two species of Parar-
hadinorhynchus, one new - - - - - - - ibs}
Edmonds, S. J., & Jamieson, B. G. M. A new genus and species of earthworm
(Megascolecidae: Oligochaeta) from South Australia - = 23
White, T.C.R. Aerial Dispersal of Adult Cardiaspina rs SORES:
Psyllidae) in South Australia - - 29
Watson, G. F., & Martin, A. A. Life history, larval morphology and pee eostips
of Australian leptodactylid frogs - - - - es
Mitchell, F. J. Studies on the Ecology of the seecAge, Lizard Amphibolurus
maculosus (Mitchell) - - - - - - - 47
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LANDSURFACE DEVELOPMENT IN THE MID NORTH OF
SOUTH AUSTRALIA
BY N. F. ALLEY*
Summary
ALLEY, N. F., 1973. Landsurface Development in the Mid North of South Australia. Trans.
R. Soc. S. Aust., 97 (1), 1-17, 28th February, 1973.
The landsurface of the Mid North has resulted from a complex tectonic and erosional history.
During the Cainozoic, faulting formed large basins in which terrestrial and marine sediments were
deposited. The earth movements also initiated two major phases of erosion.
Remnants of a landsurface established in (?) pre-Tertiary times are found high in the relief as
weathered hills and plateaux in places capped by laterite (laterite surface). A probable mid Tertiary
phase of planation is represented by silcrete capped remnants (silcrete surface). It is underlain by
deeply weathered bedrock and occurs only a few tens of metres below the laterite level. The present
broad valley floors, wide intermontane plains and coastal plains (plains and valley surface) had
developed by the Late Quaternary. Etch surfaces are found adjacent to remnants of the two earlier
surfaces.
There is considerable structural control of landforms. Resistant quartzites and sandstones form the
highest parts of the relief, as they also did during previous phases of erosion. Argillaceous strata
have been eroded to form broad valleys, which follow the regional meridional strike. Differential
erosion of these contrasting lithologies, combined with stream piracy, has led to the development of
a trellised drainage system. The River Broughton is the only stream, which has maintained its
westerly, consequent course throughout the Cainozoic, but stream captures have led to large
changes in the drainage patterns of the Light and North Para Rivers.
LANDSURFACE DEVELOPMENT IN THE MID NORTH OF SOUTH AUSTRALIA
by N. F. Avitey*
ALLEY, N. F., 1973. Lundsurface Development in the Mid North of South Australia. Trans.
R. Sov. S. Aust., 97 (1), 1-17, 28th February, 1973.
The landsurface of the Mid North has resulted from a complex teclonic and erosional
histary, During the Cainozoic, faulting formed large basins in which terrestrial and marine
sediments were deposited. The earth movements also initiated {wo major phases of erosion.
Remnants of a Jandsurface established in (?) pre-Tertiary times are found high in the
relief as weathered hills and plateaux in places capped by laterite (Jaterite surface). A
probable mid ‘Tertiary phase of planation is represented by silcrete capped remnants (silcrete
surface). It is underlain by decply weathered bedrock and ocevrs only a few tens of metres
below the laterite level. The present broad valley floors, wide intermontane plains and
coustal plains (plains and valley surface) had developed by the Lute Quaternary, Etch surfaces
are found adjacent to remnants of the two earlier surfaces,
There is considerable structural control of landforms. Resistant quartzites and sandstones
form the highest parts of the relief, as they also did during previous phases of erosion.
Argillaceous ‘strata have been eroded to form broad valleys which follow the regional
meridional strike. Differential erosion of these contrasting lithologies. combined with stream
piracy, has led to the development of a trellised drainage system. The River Broughton is
the only stream which has maintained its westerly, consequent course throughont the Camo
zoic, bul stream captures have led to Jarge changes in the drainage patterns of the Light
and North Para Rivers,
Introduction
The Mid North forms an area of moderate
relicf between the fold ridges of the Flinders
Ranges to the north and the upfaulted plateau
of the Mt. Lofty Ranges to the south (Fig. 1),
The occurrence of fluyio-lacustrine and marine
sediments on ridge tops, along river valleys and
in structural basins indicates that this Tegion
has had a complex history. Remnants of
weathered tesiduals capped by laterite or sil-
crete provide further evidence of this com-
plexity.
It is the purpose of this paper to trace the
sequence of geomorphic events which Jed to
the formation of these features and their
Preservation in the present Jandscape.
Geological Setting
The Adelaide System Rocks
The northern part of the area is underlain by
Willouran, Torrensian and Sturtian strata
(Wilson 1952; Clare 1 : 63, 360 sheet; Forbes
19651). These rocks are arranged in en echelon
folds in the northwest but parallel folds oceur
in the east (Fig. 1), South of Riverton faulting
has resulted’ in the termination and breaching
of the folds (Kapunda 1: 63, 360 sheet; Truro
1:63, 360 sheet). The Kanmantoo Group
occurs here and is underlain by a sequence of
Lower Cambrian sediments (Coats 1960*).
importance af Lithology and Faulting
The major topographic features. reflect the
contrasting rock lithologies. Between Clare
* Terrain Sciences Division. Geological Survey of Canada. Present address, Soils Division, B.C, De-
partment of Agriculture. Kelowna. British Columbla, Canada. Research carried out at the
Universily of Adelaide.
1 Korbes, B. G. (1965).—The ecology of the Clare 1: Mile area. Unpublished report, Geological
Survey of South Australia.
* Coats, R. P. (1960).—The geology of the Kapunda—Truro Military Sheets,.An explanation of the
Geological Maps. Unpublished report, Geological Survey of South Australia.
2 N. F. ALLEY
SOUTH AUSTRALIA
BARUNGA R
ANGE
—_—— ——
\ Narridy
2 Yacka Gorge
3 Spalding Gorge
4 Nantowarra Hills
--7- Approximate
dip and strike of
bedrock
|
OKm 15
[ere
Fig.
and bedrock structures are shown.
and the Murray Plains, the Gilbert Range
Quartzite underlies the major ridges whereas
dominantly argillaceous strata underlie the
valleys (Fig. 5).
Apart from the quartzite ridges, fault scarps
form the most important physiographic breaks.
The most conspicuous of these is one resulting
a Baloklava
Lr -s-
-
Vy
ty 4 h t
1 ' 4 7
1 ' ' ' ‘
' ! ‘ i r]
' 1 ‘ ' i
! 1 / t 7
© Spalding ' '
' ' 4
4 \ 4 rk 4
1 ' ‘ I
i ' t \ 1
' ' 1 rt
' ' 1 \ ro
i \ 1!
' t 1
\ ‘ a !
1 ' 1S i toy
rope 4 e +; F
4 af | r\o4 tak
U { ert rl \
too) 4 te 1 nS
Ul 1 1 : {oa \ 1 \
x 1 oo p M\er 1 \
a 5 re 2
fon
ey 1 1
‘oa ‘
rm) ‘ 1 i)
o ! 1 ‘
aw}
= F a br
1 ‘ \
& 1 ft
|
=
a !
rc) t 1
'
Undalyay - bs
1
\
a
fy
*Riverton
1. Location, points of reference and general siruciure of the Mid North, Only the major faults
from movements along the Alma Fault west of
Clare, This fault and others divide the Mid
North into a series of north-south trending
blocks (Fig. 1). The youthfulness of many
scarps and the scismicity of the area (Sutton
& White 1968) suggest that the faulting con-
tinues.
LANDSURPACE DEVELOPMENT IN 'THE MID NORTH
Cainozoic Sediments
Although Tertiary marine strata abut the
eastern and western flanks of the ranges, terres-
trial sediments of similar age occur in several
basins and valleys within the uplands (Table
vr
|). Two of the most important of the latter
are the Wakefield and Barossa Basins, Buried
beneath the sediments in these basins is a
partly dissected, weathered and faulted Jand-
surface,
TABLE 1
Terrestrial Sediments of the Mid North
Name
Unnamed
Description and occurrence
Mottled and calcareous red sandy-clays.
Found in major river valleys. intermontané
basins and coastal plains.
Age
(7) Pleistacene
tao Early Recent
Koolwnga Gravels and
siticified sands and
gravels
Rowland Flat Sands
Flavio-lacustrine sands, gravels and boulders
widespread in the Broughton caichment par-
ticularly en the castern margins of the
Wakefield Basin.
Fluvio-lacustrine sands and. gravels found
in the Barossa urea. Lignitic at depth in
the Barossa Valley,
Snowtown Sands
Interbeddéd fluvio-lacustrine
and ligniies found below
Plains.
sands, clays
the Condowte
Clinton Coal
Late Tertiary to
Quaternary
Miucene
Mincene
Middle-Upper
Measures estuarine origin, Confined to the Wakefield Ecocene
Basin,
A. PLUVIO-LACUSTRINE AND MARINE Middle-Upper Eocene beds (Harris pers.
SEDIMENTS comin, 1968). The Miocene strata are
1. WAKEFIELD Basin
The pattern of faults. demonstrate that the
busin ts of tectonic origin (Fig. 1). As it forms
the northern apex of the St. Vincent Basin,
the periodic earth movements and subsequent
sedimentation evidenced in both are necessarily
related,
Deeper Basin Sediments
Bores drilled through the Inkerman-Bala-
kKlava and Whitwarta coal measures penetrated
a considerable thickness of estuarine (Harris
pers, comm. 1968), marine and lacustrine de-
posits. (Johnson 1959; Hillwood 1960). The
sequences of the Balaklava area are complex
and interpretation of the deposits. in particular
the estuarine lignites, is uncertain, However,
recent studies refer the latter to the Middle-
Upper Eocene (the Clinton Coal Measures of
Hartis 1966), und the overlying marine beds
to the Oligo-Miacene (Hillwood 1960). ‘The
northern and eastern limits of the marine beds
lic along the Bowmans and Mt. ‘Templeton
Fuults. At Snowtown the marine deposits are
absent but interbedded carbonaceous clays,
sills und sands of Miocene age overlie the
lacustrine and terminate in the south along the
Balaklava Fault, Because of their stratigraphic
relationship to the Early Tertiary beds and
their dominantly sandy lithology, the Miocene
strata are separaled from the former and here
named the Snawrown Sands, The type section
is from stratigraphic borehole 4A on the
northern edge of Snowtown and the strati-
graphic relationships are shown in Figure 6.
‘The mid Tertiary marine sequences and the
Snowtuwn Sands are separated from each other
by «a horst, known as the Nantawatra Hills,
which resulted from uplift along the Bowmans
and Balaklava Faults {Fig. 1). The absence
of the Oligo-Miocene sediments from the sur-
face of the horst and their occurrence at the
foot of the bounding fault scarps on the north
and south suggests that the faulting predates
the Middle Tertiary.
Marginal Sediments
Numerous deposits of highly siliceous sands
and gravels occur along the fringes of the
Wakefield Basin. Small outcrops abut the
eastern front of the Barunga Range but Jarger
deposits occur in the valley of the River
Broughton on ihe eastern edgé of the basin.
N. FL ALLEY
&
4 Gulnare
Fluvios (ucustrine deposits. _-
Surface exposures of the ~ 4)
Koolunga Gravels :
Uplands,
OKm 5
——
‘Alma Fault
2. The distribution of the Koolunga Gravels in the middle and upper Broughton drainage basin.
The dips. of the faults in the Mt, Gregory Range are unknown,
The most prominen| crop out near Koolinga.
Yacka and Spalding (Fig, 2). These deposits
are here named the Koolungu Gravels and the
lype section is located at White Cliffs, 4 km
east of Koolunga in the south bank of the
Broughton River (Fig. 10).
The western edge of the Koulunga Gravels
is marked by a low rise which may represent
the northern extension of the Owen Fault
Scarp, for bore logs indicate that they continue
al depth further westwards. At White Cliffs
the gravels overlie an. itregular surface eroded
across Precambrian bedrock (Fig. 10). Move-
ment along a fault zone in the latter has led
toa J m dislocation of the overlying deposits.
The best exposure of the beds occurs in the
Yucka Valley where deltaic sands and gravels
containing boulders of silcrete overlie gently
dipping Precambrian sandstone. These deposits
can be traced across the ridge marking the
western flank of the valley, but the Alma Fault
scarp separates them from the Koolunga
sequences (Fig, 2). Because of their similar
pebble lithologies, in particular the presence
of silerete in both, the Yacka and Koolunga
beds were probably deposited contempor-
angously and have since been displaced by
faulting with the downthrow to the west.
Whether the highly siliceous gravels south
and cust of Spalding were deposited in the
same lake is a matter for conjecture, for they
stand 100 m higher than those at Yacka (Fig
2). The difference in height between the two
deposits may reflect a fallmg lake level or fault-
ing, or the Spalding gravels may not be corre-
lative with the Yacka heds and instead be en-
tirely fluviatile, However, the presence of
xilerele houlders in the Spalding gravels indi-
cutes that they may have been deposited con-
temporaneously with the Koolunga Gravels.
Although no fossils were recovered from the
Koolunga Gravels, there 1s evidence to suggest
that they post-date the mid Tertiary, At White
Cliffs and Yacka the gravels contain boulders
of silcrete. a rock assigned lo a probable Middle
IT ANDSURFACE DEVELOPMENT IN THE MID NORTH 5
Tertiary weathering phase (see below). North
and west of Yacka, outcrops of the gravels
overlie an etch surface related to the erosion
nf the silcrete surface (Fig, 3). Thus, the
Koolunga Gravels do not represent a shoreline
facies of the Snowtown Sands but were de-
posited at different times in a Jake occupying
the same basin.
Silerete &
~koolunga
a
Gravels
(exhumed }
Fig. 3. The stratigraphic relationship between
the Koolunga Gravels und the silcrete
surface in the Yacka area, Only the sur-
face exposures of the gravels are shown.
2, Bakossa Basin
These include the fluvio-lacustrine sediments
deposited east of the Para Fault in the Barossa
and Stockport-Freeling areas (Pig. 4).
The strugture of the Barossa Valley is not
completely Known. Tt has been described as a
graben (Hossfeld 1949), a Fault angle valley
(Chuge 1955), and an asymmetrical synctine
with q fault or flexure along the southeastern
flank (Dalgarno 19613; Olliver 1962). Never-
theless, the nature of the Tertiary strata ¢on-
tained by the valley is well documented (for a
review of this work see Dalgamno 1961, and
Harris & Olver 1965). West of Rowland Flat
a basal deltaic unit is overlain by a sequence
of sands and gravels characterized by medium
und Jarge scale deltaic bedding (the Rowland
Flat Sands Conglomeratic Lens and the Row-
hind Flat Sands of Dalgarno 1961), whereas
lignitie sands have been encountered at a
depth of 30 ta 100 m during boring in the
Rowland Flat-Tanunda plain (Chugg 1955),
As there is no stratigraphic evidence for
scpurating the conglomeratic lens from the
overlying sands, it is proposed that it be in-
cluded as part of the Rowland Flat Sunds.
Scattered outcrops of the sands extend west-
wards across the Kitchener Fault to the Gawler
area, and northwards to Stockport (Fig. 4).
They were probably deposited contem-
porancously with the Rowland Flat Sands as
the beds occur within the same range of eleva-
tion (Fig. 4). Moreover, the Lyndoch-Sandy
Creek deposits ure characterized by the same
mineral assemblage as those at Rowland Flat
and are thus regarded as the westerly extension
of the Barossa sequences (Olliver & Weir
1967).
Although the Rawland Flat Sands. have been
assigned a variety of ages, recent palynological
evidence indicates « Miocene age, making them
time correlative with the deposits northeast
of Gawler (Haris pers. comm. 1969).
B. ALLUVIAL DEPOSITS
Red alluvium covers most of the low-lying
iireas, varying from a Tew metres in minor
creeks to 36 m in depth at Balaklava. In the
Barossa Valley and near Red Ranks, bores
show that the sandy clays exceed 30 m in
thickness. Although the alluvium is 15 m deep
in the Yackamoorundie Valley downstream
from Georgetown, it rarely exceeds 6 mm in the
upper reaches of other main Valleys, In most
areas the alluvium is mottled and calcareous,
Suggesting some measure of weathering since
its deposition.
The alluvial clays are interpreted as a
Pleistocene deposit in the Port Wakeficld area
(Johnson 1959; Hillwood 19640; Horwitz 1961)
7 Dalgarno, C: R. (1961}—Geology of the Barossa Valley. Unpublished M.Sc. Thesis. Universily of
Adelaide.
6 N. F. ALLEY
Sandy Creek
"215 Spot Height imetres! Q cnt
266
Para Fault
Fig.
and the Burossa Walley (Campana 1955;
Chugg 1955). On the basis of radiocarbon dat-
ing, the red clays. north of Adelaide have been
assigned to the Late Pleistocene (Williams
1969}, Mottled clays (the Hindmarsh Clay of
Firman, pers. camm, 1968), overlying a Plio-
cene marine bed at Red Banks are believed to
be of Pleistocene age.
On this basis the heavily calcified red clays
in the Mid North have been placed tentatively
fee
5 lyndoch, SS
sKAPUNDA
-—— Moin uplands
=| Rewland Flat
: Sands (sic. exo.)
4. The distribution of the Rowland Flat Sands in the Barossa-Freeling area. The differences in
height of the deposits in the latitude of Freeling may be related to post Miocene earth move-
ments,
in the Pleistocene to Early Recent though they
may vary in their precise age or age-range from
locality to localily.
C, WEATHERED LAND SURFACES
BURIED BENEATH THE CAINOZOIC
SEDIMENTS
In the Snowtown area only a few bores
penetrate to the Precambrian but these reveal
that the basin floor is irregular and weathered.
LANDSURFACE DEVELOPMENT IN THE MID NORTH 7
At a depth of 69 m, stratigraphic bore No. 4A
located at Snowtown, penetrated 27 m of
highly weathered sandstone, Three kilometres
north of Snowtown unweathered bedrock
occurs at 49 m, but Icss than 1.5 ko east of
this location the basin floor deepens to 111 m..
{ f
OMintate f
a
-
ep
©
‘The weathering of the basement rocks probably
predates. the deposition of the overlying Ter-
tiary sequences for bore logs indicate that the
latter are essentially unweathered,
The nature of the buried surface beneath the
Rowland Flat Sands is better known. Bore logs
Q Merildin
S lerete leyrmeants
¥e8 +566 Spot height (r
=
OWotervale
a Gilbert Ronge Quartzite
Weathered residual) and etch
plains related to the
laterite level
=— Loierite level
ea
Fig. 5. Remnants of the laterite and silcrete surfaces near Merildin. The Gilbert Range Quarizite
underlies the major ridges, while dominantly argilliceous strata crop out in the valleys.”
+ WN, F. ALLEY
indicate that the basement rocks are weathered
in the Barossa Valley (Chuge 1955). At Row-
land Flat the unconformity surface has a local
telief of up ta 15 m (Olliver & Weir 1967)
and the Rowland Flat Sands overlie both fresh
and weathered bedrock (Fig. Ll). At ©. R,
Hueppau!’s quarry, on the north bank of the
North Para River, Torrensian bedrock is
weathered and highly ferruginized. Between
Sandy Creek and Warpoo Siding, highly
kuolinized mica schist underlies the sands.
South and west of Frecling the sands were de-
posited on subhorizontal bedrock surfaces,
though only small remnants of the Tertiary
sequences remain. Thus it appears that the
Rowland Flat Sands were deposited on a partly
dissecied erosion surface underlain by highly
weathered and ferruginized bedrock.
Surfaces uf Degradation and Deposition
Investigations reveal that remnants of three
major surfaces of erasion and deposition occur
in the Mid North. The highest and oldest of
these is capped discontinuously with laterite.
Below this is another erosional surface which
supports a thick crust of silcrete, The youngest
surface ls largely depositional and the alluvial
clays underling it ure heavily calcified, Etch
plains of considerable extent developed as a
result of the dissection of the laterite and sil-
crete surfaces,
Larertte Surface
Apart from the prominent ridges underlain
by quartzite and sandstone, remnants of the
laterite surface form the highest parts of the
lajdscape. Where, the laterite truncates thick
Precambrian argillaccous strata, slopes vary
belween 1° and 2° (Fig, 12), However, near
the quartzite ridges remnants of pediments
exhibiting slopes of up to 7° are found, Such
residuals are common in the ranges euse of
Clare (Fig. 5) and Riverton, Remnants of the
laterite surface are more numerous at the
streum) Heads. Nevertheless, by virtue of their
resistant capping they alsa fonm prominent. hills
standing nearly 100 m above valley floors.
yome distance from the ridges (Fig. 13).
The laterite capping, which consists essen-
tially of angular quartz set in a matrix of tron
oxides, is underlain by heavily weathered and
locally kaolinitic bedrock (Fig. 14>. Only the
massive quartzites and sandstones remain uny-
altered and these stand 30-180 m ubove the
general surface level) A zone of intense
weathering often lies at Ihe head of the pedi-
{nent Férnnants Which slope away from these
scarps wheras the Jaterite crust is consistently
thickest on the lower slopes.
The etching away of the weathered strata
from beneath the laterite has let to the
exposure of broad bedrock surfaces, It is
usually the case that these etch surfaces lie
15-30 m below the laterite surface and
separated from it by seep scarps. Because of
strong lithological control during weathering
and subsequent erosion, the etch surfaces are
often irregular with argillaceous strata forming
the vallevs, and sandstones the ridges (Fiz.
13),
Silerete Surface
Evidence of a former erosion surface capped
by silcrete and consistently lower than the
height of the faterite is found throughout the
Mid North (Fig. 15). It reaches its optimum
development around the northeastern fringe
of the Wakefield Basin (Fig, 6), Here slopes
can be less than 1°” whereas slopes as great
a8 4° have heen encountered elsewhere. Like
the laterite, the silerete surface truncates rocks
of ull types, except the thick quartzites, and
1s Underdain by some 15 m of intensely ksolin-
ized bedrock. North of Narridy the weathering
reaches a depth of 30 m, bul this is probably
duc to the porosity of the underlying sandstonc.
Fertuginuus hurtzuns are common in the
weathered zone developed across argillaccous
strata (Fig, 7),
‘The nature of the silerete crust is variable
(Figs. 7, 16). Higher parts of the surface are
weakly silicified or suppart a thin capping of
consolidated regolith. Silcrete lower in the
relief is generally thicker und finer grained, In
many areas where the silcrete has heen broken
up. a thick lag of boulders remains as a pro-
tective veneer over the weathered bedrock,
An extensive etch plain developed following
dissection of the silerete surface, particularly
where the main streams cut deep gorges
through the silercte level. Thus prominent bed-
rock benches occur slong the Rocky Rivet
downstream from Gladstone and the Yacka-
moorundie Creek west of Gulnare. This sur-
face is often capped by a silicificd congtom-
erle containing Lragments of silcrete derived
from adjacent silcrete remnants (Fig, 17),
Because the conglomerate has all the charac-
|_ANDSURFACE DEVELOPMENT IN THE MID: NORTH 9
N
4
4
7
4
i]
qt
=
4
1
oBurra _ 4
|
Murray
Laterite Bid:
ans
remndnts
dominate
Largely silcrete remnants
oy
Areas farming
probable Miocene lake basins
~
~
3
o
-~
s Gkm
| ce |
<<
= cr 2 f
as DGawler-—-E= ——
ras Leterjte
Nantawarro Hills
A a i Silcrete S
a Lote Quaternary alluvium
~*< = =a = ee
S 3 2
\ = Weathered Pre £
~ ri ~~, “
ee
dn
Balaklava Fault
Fig, 6, The distribution of laterite and silcrete remnants in the Mid North, and the stratigraphic re-
lationship between the surfaces bordering the Wakefield Basin. The diagrammatic section
(not to scale) has been constructed from bare logs. From the surface, 1$ m of plains and valley
alluvium are underlain by 15 m of silts and clays, 23 m of Miocene Snowtown Sands and 35
m of Middle-Upper Eocene Clinton Coal Measures (E). Both the latter units vary consider-
ably in thickness, The Mt, Templeton Fault occurs in the breached part of the section.
(a N. F_ ALLEY
Raolinized
sondstone
,
f
‘gin oF, tren oxides
a | fr
3. West of Kupunde
Fig, 7. Variations in the silcrete crust and its
underlying weathering profile in the Mid
North,
Section 1: Zone A consists of pisolitic
silcrete; B laminar sitcrete; C is a com:
plex zone grading downwards from sili-
cified sandstone to very resistant por-
celanitic silcrete.
Section 2: Zone A, massive crypto-
crystalline silerete; B, weakly silicified
sandstone; C, very weathered rock con-
tuining abundant angular fragments of
quartz.
Section 3: Zone A, massive crypto-
crysialline silercte containing numcrous
rounded quartz pebbles near its base.
teristics of a sedimentary rock and uncon-
formably overlies fresh Precambrian at
numerous sites, it is not interpreted as a second
period of silcrete formation and is referred to
as silicified sands and gravels. On the basis of
its relationship to the etch surface these sedi-
menis were probably deposited in valley
bottoms contemporancously with the Koolunga
Gravels.
Relative Ages of the Laterite and Silcrete
The laterite surface forms the flat upland of
the Nantawarra Hills (Fig. 6), To the ease
and north of the hills, the surface has been
downfaulted and the Middle Tertiary sedi-
ments occur at the base of the fault scarps
suggesting that the laterite surface in this areca
may be Early Tertiary or older, In this case,
the surface buried beneath the Middlc-Wpper
Eocene lignites in the Snowtown region could
be the equivalent of the laterite surface, On
the other hand, if the lateritized surface un-
derlying the Rowland Plat Sands is correlative
with the laterite surface, the Jatter must have
persisted in the Barossa area until the mid
Tertiary when earth movements led to its dis-
section and partial burial.
ii is difficult to assign an age range to the
silerete surface, although mapping reveals that
it is consistently Jower than the laterite surface
even where they truncate the same strata (Fig.
8), Thus the difference in their heights is due
to crosion rather than Jithology, and it can be
inferred that the silerete is younger than the
laterite surface. However, the optimum de-
velopment of the silcrete surface occurs around
the fringes of the Wakefield Busin, There is
evidence lo suggest that the early ‘Tertiary
Fig. 8. The altitudinal relationship between the
laterite and silcrete: surfaces west of
Georgetown. The bedrock strike is
approximately north-south, ard both
surfaces are eroded across the same
strata,
LANDSURFACE DEVELOPMENT IN THE MID NORTH It
esluatine swamps and then the Miocene lake
which occupied this basin may have formed
the base level af erosion for the silcrete sur-
face in the surroundiig area, First, the surface
slopes vently towards the basin. Second, the
Eocene estuarine and the Miocene Jacustrine
sequences are generally fine, suggesting that
deposition Was cantemporaneous with the ero-
sion of a plain of low relief which is consistent
with the gentle slopes observed on the silcrete
remnants, Third. no silcrete is recorded from
the borcholes drilled through the Tertiary
secliments,
Present Valley Floors and Plains
The must extensive surface of ageradation in
the study area is the Condowie Plain (Fig.
1), Near Snowtown the plain measures almost
4 km from east to west, whereas in the
noflbern part near Redhill the basin is Jess
than § km wide. The Condowie Pliin con-
tinkes southwards and gradually merges with
the Adelaide coastal plains south of Baluklava.
In the latitude of Snowtown the surface of the
plain is gently undulating but asymmetrical
wilh a long. fall from the Alma Fault scarp
on the east. No streams traverse the plains
excep! along the northern fringes: Where the
Broughton and its two lurge tributaries, Rocky
River and Yackamoorundie Creek, are incised
up to 9 m in the red alluvium.
Two other intermontane basing oceur: the
Barossa Valley and athe Yackamoorundie
Valley south of Georgetown, the later con-
twining an alluvial fill 54 m in depth. The
surfice of the Barossa is similar to the Con
dowie Plain. displaying east-west asymmetry.
Broad alluvial Fans front the Barossa Ranges
near Rowland Plat, gradually giving way to a
gently sloping surface further north. Along the
westerh edge of the basin the plain extends
across aulcrops of the Rowland Flat Sands as
a lightly ferruginized erosional surface.
Age of the Plains and Valley Supfice
Vhroughout the Mid Novth, the red cal-
carcous clays are the most common deposits
underlying the surface. This alluvium has been
assigned a Pleistocene to Recent age (see
above), and thus the surface must be approxi-
mately the sume age, In the Barossa where it
(runciles the Middle ‘Tertiary lake sediments,
portion of the surface may belong to the Late
‘Terriary
Development of the Drainage System
The Mid North is drained by two sets of
river systems: the Broughton, flowing to Spén-
cer Gulf, and the Wakefield, Light ani North
Para Rivers. to St. Vincent Gulf, The history
of change in this drainage pattern can be
determined by an examination of the nature of
the Tertiary sediments and landsurfaces.
A. INFLUENCE OF LITHOLOGY AND
STRUCTURE
The perfection of lithologies) adaptation of
the drainage is a feature of the area. In broad
view lithology determined the direction of
drainage on the two duricrusted surfaces, Once
the weathered mantle was removed from these
surfaces. intricate irregularities in lithology
controlled the course of erosion. Consequently,
the present drainage exhibits 4 trellised pat-
tern, the long subsequent (strike) streams
flowing in valleys underlain by argillites while
the resistant sandstones and quastzites remain
as the fidges. The north-south trending prat-
fern of the major streams results from = the
meridional strike of the fold axes (Fig. 11.
B. CHANGES IN THE PATTERN DURING
THE CAINGZOIC
The long meridional trending reaches of the
strcams have heen interpreted as the dismem-
bered drainage of a Tertiary peneplain (How-
chin 1933; Hossfeld 1935; Langford-Smith
1949), Such a postulate may only be applied
to the St. Vincent Guif drainage.
1. Tie River BroucHTon
Based on the evidence of elbows of capture,
abandoned valleys containing riverine graveis
and increased gradients in the gorge sectors of
the newly formed valleys, Langford-Smith
(1949) demonstrated that stream piracy
occurved amongst minor tributaries in the
Jumestown area. In view ol the elbows and
gorges slong the Broughton, he applied the
theory to the whole drainage basi, He pro-
posxd that the river captured a system of
southerly flowing streams from ihe west sub-
sequent upon uplift of a Tertiary Jundsurface.
Faulting aided local ponding of the stream
and hastened the speed of capture.
However, investigations cannot substantiate
the theory. Only two gorges occu’ along the
Broughton, ohne west of Yacka and the other
west of Spalding, No knickpoini is found in the
Yacka reach while those in the Spalding gorge
fesulk [rom lithology or structure. There is no
2 N, F. ALLEY
evidence of a former south flowing stream
system, the development of which was pro-
hibited by the geologic structure of the arca
and the resistance of the Gilbert Range Quart-
zite, Upstream from Yacka the drainage pat-
tem may have been in existence since Early
Tertiary times, for remnants of the laterite
surface slope towards major streams whereas
their heights decrease down-valley in the direc-
tion of present drainage. Many of the Caino-
zoic gravels, some of them lacustrine, occur
both on intervening ridges and in several
gorges, instead of only in the broad strike
valleys, As uplift of the Tertiary landsurface
occurred along N-S trending faults, there
should have been no local impounding of south
flowing rivers. Moreover, the disposition of the
Cainozoic sequences in the Yacka-Spalding
area suggests faulting after depositton, not be-
fore. Finally, if stream piracy has occurred, it
is the Broughton system which is being cap-
tured not the reverse (Fig. 9}.
The middle and upper reaches of the
Broughton have remained essentially unmodi-
fied and it is only downstream from Yacka
é
*Menldin
. Stream capture of the upper Farrell
Creek near Merildin. The nature of the
barbed drainage within the enclosed sec-
iron and the pronounced elbow of the
Wakefield downstream from Merildin
indicates. that the pirate stream was the
River Wakefield.
that the evidence suports changes in the put-
tern.
Focene. lignites extend at depth to within
a few kilometres of Koolunga where the
Broughton turns north-westwards to flow
across the northern perimeter of the Condowie
Plain. If the upper reaches have altered little
since the Early Tertiary, it is logical that the
river made its way to the present St, Vincent
Gulf area through the tectonic yalley which
came into existence at this time (sce aboye),
lt is probable that uplift of the Nantawarra
horst in the mid Tertiary blucked the valley,
leading to the formation of a lake, A larger
lake resulting from reactivation of upwarping
in the Late Tertiary or Quaternary either over-
flowed the Baruhga Range or was captured
from the west (or both) and the Broughton
fiowed into Spencer Gulf.
2. St, VINCENT GULF DRAINAGE
A striking feature of the drainage pattern
of these rivers. is the absence of major south
bank tributaries (Fig. 1), An eaamination of
the Cainozoic sediments and the Tertiary land-
surfaces reveals that only stream piracy can
account for the anomaly.
The long quartzite ridge east of Tarlee was
a major drainage divide in Early Tertiary
times. All laterite remnants slope away frem it
as far north as Manoora and south as Green-
ock, Erosion of this surface by separate drain-
uge systems is indicated by large diiferences
in the height of residuals on opposite flanks
of lhe ridge. The river east of the ridge occu-
pied the valley of the upper River Light und
flowed southwards along the present Barossa
Valley area whereas the other approximutely
paralleled the course of the Gilbert River.
However, there ts no evidence to suggest that
the River Wakefield had yet been established.
Consequent upon the mid Tertiary earth
movements were the dissection of the laterite
surface and the formation of a Jake in which
the Rowland Flat Sands were deposited. The
two south flowing streams probably enterudd
the lake at the northern edge of the Barossa
Valley and near Stockport. This is borne owt in
these areas by the coarseness of the gravels,
the palacocurrents and the mineral assemblage
(Dalgarno 1961, Alley 19694).
In Late Cainozoic times the majar stream
east of the divide was captured from the west
LANDSURFACE DEVELOPMENT IN THE MID NORTH 13
by two separate streams, one forming the
present lower reaches of the River Light now
confluent with the Gilbert River, and the
other, the North Para River. Stream piracy is
supported by several lines of evidence, First,
prominent elbows occur on the rivers where
they turn westwards to flow across the strike
of the bedrock (Fig. 1). Second, an abandoned
valley due south of Rowland Fiat and infilled
with Tertiary sediments may represent the
former channel of the beheaded stream. Third,
hoth portions of the newly formed streams
have cut deep charinels into the Rowland Flat
Sands, suggesting that the excavation post-dates
the mid Tertiary,
Although a prominent elbow occurs en the
River Wakefield at Undalya, where the river
turns into a gorge and flows across the strike
nothing other than the lack of major south
bank tributaries to suggest that capture may
have taken place,
Landscape History
Remnants of three major Jandsurfaces have
been described and discussed (Table 2). Since
the dissection of the Jaterite surface, differen-
tial erosion has led to increasing relief. Where
the ridges formerly stood 90 to 180 m above
the level of the laterite surface, they now pro~
ject more than 300 m above valley floors.
Early Tertiary earth movements in and
around the Wakefield Basin probably led to
the disruption of the laterite surface in the
Broughton drainage system and the establish-
ment of a new phase of erosion during which
of extremely resistant
TABLE 2
Surfaces in the Mid North
quartzites, there ts the silcrete surface developed. Broad etch sur-
Surface
Plains and valley
Description and origin
Major valley floors, intermontane basins
and coastal plains, Narrow. erosional pied-
mont zone fringing these areas, Mainly
aggradational.
Unweathered bedrock surfaces ‘standing
below and adjacent to the silcrete surface
in the Broughton catchment. Developed
subsequent te removal of weathered zene
beneath the silérete, Buried by lacusitine
gravels in some localities bul exhumed jn
others,
Probable age
Late Tertiary (7?) to
Early Recent
Initiated in the Late
Cainzaic
Silcrete
Silicified mesas and valley sides occurring
throughout the Broughton catchment. fso-
lated remnants near Kapunda, Freeling and
Owen. Resulted from erosion and weather-
ing.
High unweathered bedrock benches devel-
oped below and adjacent to the laterife sur-
face as a result of removal of the weathered
mantle from below the laterite.
Laterite
Broad plateaux and mesas capped by later-
ite and, apart from quartzite ridges, form-
ing the highest parts of the relief. Resulted
from erosion and weathering.
Mid Tertiary
Initiated in the Early
Tertiary
{7) Early to pre
Tertiary. Persisted to
mid Tertiary in the
Barossa area.
faces resulted from the removal of the deeply
weathered muntle much of which was de-
posited lower in the relief as. the Middle-Upper
Eocene sediments, and later in the Miocene
as the Snowtown Sands. The laterite surface
persisted in the Barossa area until the mid
Tertiary when renewal of the faulting led to
its dissection, The earth movements probably
dammed streams, leading to the formation of
a basin in which the Rowland Flat Sands ac-
4 Alley, N. F. (1969).—The Cainozoic history af the Mid North of South Australia. Unpublished
M.A. Thesis, University of Adelaide.
14 N
cumulated, As these sands and the Snowtown
Sands are contemporaneous but deposited in
separate tectonic basins, it is logical to argue
that the earth movements were widespread
during the mid Tertiary,
There is evidence to suggest that the fault-
ing continued, In the narth, earth movements
account for the dissection of the silcrete sur
face, the development of broad etch plains and
the deposition of the Koolunga Gravels con-
temporancously with the silicified sands and
gravels. Recent earth movements were respon-
sible for the dislacation of the Koolunga
Gravels at White Cliffs and in the vicinity of
Yacka. Deposition of the plains and valley
alluvium and the alluviul fans abutting the
Alma and Kulpara Fault scarps may also have
heen initiated during this period.
If the buried surfaces beneath the Tertiary
sediments in ihe Wakefield Basin and the
RB ALLEY
Barossa Valley are analagous to the Jaterite
surface, it has suffered faulting or warping in
the order of 300 and 180 m respectively in
these areas.
Acknowledgements
The thanks of the writer are due to: the
South Australian Department of Mines for
making bore Jog data and unpublished reports
available; Mr. W. K. Harris, 8, Aust. Depart-
ment of Mines, for his advice on the Tertiary
stratigraphy; Mr. A. R. Milnes, Department of
Geology, University of Adelaide, for his
guidance in the analysis of the duricrust pro-
files; Mr. M. J, C. Walker, University of Edin-
burgh, for his critical reading of the text; and
Dr, C. R. Twidale, Department of Geography,
University of Adelaide, for his valuable com-
ments in the ficld and on this paper.
References
Camerana, B. (1955) —Lhe geology of the Gawler
Military Sheet. Rep. Invest. geal Surv. §.
Ast, 4, 1-22.
Cuucs, R. — (1955)—The hydrology of the
Rarossy Valley, Refi. Invest. geol. Surv. 8.
Aust, 2, 1-77,
Coats, R. P. (1959)—Trure map sheet, Geolo-
gical Ailaus. of South Anstralia, 1763, 360
series, (Geol, Surv. 8, Aust.: Adelaide.)
Dickmson, 8. B. & Coats, R. PB. (1957)—
Kapunda map sheet, Geological Atlas of
South Australia, 1:63, 360 scrics. (Geal.
Surv. S. Aust.: Adelaide.)
Fortis, B. G. (1964)-—Clare imap sheet, Geo-
logical Atlas of South Australia, 1:63, 360
series. (Geol, Surv, S, Aust: Adelatde.)
Harris, W. K. (1966) —New and redefined names
in South Australian Lower Tertiury strut
graphy. Quart. geol. Notes, geol, Sun §
aust. No. 20.
Harais, W. K. & Ouniver, J. G. (1965),--The
age of the Tertiary sands at Rowland Fiat,
Burossa Valley, Quart, geal. Notes, geal.
Sarv. 8, Aust. Na. $3,
Hitt woop, E. R. (1960)—Inkerman-Balaklava
coalfield, Min, Rev. Adelaide 112, 26-46,
Hoawitz, R. C. (1961).—The geology of the
Wakefield Military Sheet. Rep. Invest. geal.
Sary. §, Aust. 1B, 1-32,
Hosseeep, P. S. (1935)—The geology of part
of the north Mt. Tafly Ranges, Tras. R. Soc.
S. Aust. 59, 16-67.
Fie 1, At White Cliffs, 4 km cast af Koolunga, Late Cainozoic (7) Muvio-lacustrine sands and gravels
containing boulders of silerete have been folded: The monoclinal fold (M) indicates that 4 to
5 m of displacement has occurred. The folding is of the posthumous type and hax resulted
from movernents in the underlying contorted Precambrian phyllitic shalcs which upper in
the lower left (P)-.
Fig. |!, The irregular unconformity surface between the subhorizontally bedded, Miocene Rowland
Flat Sands. and steeply dipping Precambrian schists in a quarry west of Rowland Flu,
Fig, 12, A few kilometres west of Georgetown temnants of the laterite surface occur, Here the surface
slopes gently away from an unweathered core of Precambrian rocks known us Mt. Herbert
(right), The laterite crust is underlain by 2 to 25 m of kaolinized shales some of which
can be seen forming the white areas on the flanks of the two residuals,
Fig.
13. ML Allen. a lateritic, weathered residual eroded across green silty shales in the snout of a
syjicline & km due north of Kapunda. The arcuate, low ridge covered by gums and almost sur-
rounding Mt. Allen consists of unweathered Marinoan quartzite (A,B.C.7}. Differential ere-
sion of adjacent weathered strata, has. accentuated the prominence of the ridge and resulted in
an irtegolar etch surtace.
1s
MID NORTH
IN THE
| ANDSURFACE DEVELOPMENT
——
— | =
ALLEY
N. F.
16
NAYS
<a
ae
?
:
LANDSURFACE DEVELOPMENT IN THE MID NORTH 7
HossFELb, P, S. (1949).—The significance of the
occurrence of fossil fruits in the Barossa
sekungsfeld, Trans. R. Soc. S. Aust, 72, 252-
258,
Howcuin, W. (1933).—The dead rivers of South
Australia, Part IT. The eastern group, Trans.
R. Soc, S. Aust, 37, 1-41.
JauNson, W. (1959).—Coal exploration, northern
end of the St. Vincent Basin. Adin, Rev-
Adelaide 110, 134-144.
Jounson, W. (1964).—8urra map sheet,’ Geo-
logical Atlas of South Australia. 1763, 360
series. (Geol. Surv. S$. Aust.: Adelaide.)
Lancrorp-SmMitH, TT. (1949),—The geomor-
phology of the County Victoria. South Aus-
tralia. Trans. R. Soc, S. Aust, 72, 259-275.
Outiver, J. G. (1962).—Test boring—Rowland
Flat Sand deposit. Min. Rev. Adelaide 121,
144-150.
Ox.iver, J. G. & Wrir, L. J. (1967).—The con-
Struction sand industry in the Adelaide met-
ropolitan area, Rep, Invest. geol. Surv. 8.
Aust. 30. 1-106.
Sutton, D, J, & Wnite, R. E. (1968).—The
seismicity of South Australia, J. geol, Soc.
Aust. 18, 25-32.
WiritaMs, G, E. (1969).—Glacial age of pied-
mont alluvial deposits in the Adelaide area,
Sauth Anstralia. Aust. J. Sei, 32, 257.
Witson, A. F. (1952)—The Adelaide System as
developed in the Riverton-Clare region.
northern Mt. Lofty Ranges, South Australia.
Trans. R. Soc, S. Aust, 75, 131-149.
Fig. 14. Highly weathered Sturtian slates and gritty shales exposed beneath the laterite surface in
Bartsch’s quarry west of Kapunda. The rock is locally kaolinitic, the Iatler being closely re-
lated to bedding planes and joints. Hcight of the section is ahout 15 m.
Fig. 15. Silcrete capping the crest of an erosional scarp several kilometres north of Narridy. In places
the underlying porous sandstone is weathered &) 4 depth of 30 m or more (Fig, 7. Section 1).
An etch surface occurs to the left and lower than the silerete,
Fig, 16. Pisolitic silerete capping portion of the scarp north of Narridy (see Fig. 15). The larger con-
cretions consist entirely of slightly ferruginous silerete, but smaller nodules contain porcelan-
ous kaolin, small rounded grains of silica and a few minor pockets of iron oxides. The piso-
liths are set in a matrix of slightly silicified sandstone. Pen provides scale.
Fig. 17,
Boulders of the silicified sands and gravels (right) scattered over an etch surface developed
adjacent to the silerete surface Cleft), in the valley of the Rocky River several kilometres
south of Gladstone, The gravels unconlormably overlie steeply dipping unweathered Torren-
sian slates and shales.
AUSTRALIAN ACANTHOCEPHALA, NO. 14. ON TWO SPECIES OF
PARARHADINORHYNCHUS, ONE NEW
BY S. J. EDMONDS*
Summary
EDMONDS, S. J., 1973. Australian Acanthocephala, No. 14. On Two Species of
Pararhadinorhynchus, one new. Trans. R. Soc. S. Aust. 97 (1), 19-21, 28th February. 1973.
A new species of Acanthocephala, Pararhadinorhynchus coorongensis, is described from the fish
Aldrichetta forsteri (Cuvier & Valenciennes) from South Australia.
AUSTRALIAN ACANTHOCEPHALA, No, 14. ON TWO SPECIES OF
PARARHADINORHYNCHUS, ONE NEW
by S. J, EoMonps*
Summaty
Epmonps, §. J.. 1973. Australian Acanthocephala, No. 14. On Two Species of Pararhadineor-
Avachus, one new. Trans. R, Soc. §. Aust. 97 (1), 19-24, 28th February, 1973.
A new species of Acanthocephala, Pardrhadinorhynchus coorongensis, is described from
the fish Aldrichetta forsteri (Cuvier & Valenciennes) from South Australia.
Introduction
Johnston & Edmonds (1947) erected a new
venus Pararhadinorhynchus based on P.. mugi-
fis, a parasite from Mugil cephalus Linnaeus.
On several occasions during the last ten years,
a second species of the same genus has been
found in South Australia in the mullet Aldri-
chetta (=Agonestomus) forsteri (Cuvier &
Valenciennes), The new species is described
here and the re-checked dimensions of the
introvert and egg of P. mugilis are given. How
Specific the two parasites are in their distribu-
tion is not clear because collectors have some-
limes confused the two hosts. The identifica-
tions of the fishes carrying the parasites are
based on the descriptions given by Scott
(1942).
Pararhadinorhynchys coorongensis n.sp,
FIGS. 1-5
Host: Aldrichetia forsreri (Cuvier & Valen-
ciennes).
Localities; (1) Coorong, S.A.: coll. J. Harris,
1962. (2) Port Willunga, S.A.; coll. T. H.
Johnston, 1927: (H.C, 1055). (3) Fish
from Adelaide Fish Market (class material.
coll. &. J, Edmonds, 1969, 1971). (4) Pish
from Adelaide Fish Market: coll. H. Man-
ter, 1967,
Type specimens (male and female): Aus-
tralian Museum, Sydney.
Description: Size moderate; shape cylindrical
to sub-spindle-like, slightly broader in anterior
third or quarter. Posterior region may taper
slightly, Posterior extremity sometimes
rounded or slightly swollen,
Trunk of female 9-15 mm long x 0.45-
0.55 mm. Trunk of male 7-11 mm x 0,40—
0.70 mm. Trunk without spines.
Introvert. cylindrical to club-shaped; the part
bearing hooks 0.51—0.62 mm long in female
and 0,50-0.58 mm in male. Maximum width
0.15-0.22 mm in female and 0,12-0.23 mm in
male, Fourteen to sixteen, usually 16, longi-
tudinal rows of 8-10, usually 9, hooks per
row (Fig. 2). Short unarmed neck (0.15 mm
long) in some specimens. Introvert sheath
double walled, 0,6-0.8 mm long x 0.18-0.26
mm wide. Lemnisci about twice as long as
sheath and usually slightly swollen posteriorly,
Position of brain not known,
Female complex relatively long, 2,8-3.4
mm. Female aperture terminal. Ripe eggs,
possessing prolongations of middle shell,
.042-0.046 min x 0.008-0,010 mm.
Testes ellipsoidal to subglabular and placed
in tandem; anterior one 0.45-1.05 mm x 0.35—
0.45 mm and posterior one 0.50-0.91 mm x
0.34-0.43. Cement glands, two, Jong and
slender, sometimes constricted at some points.
Male aperture terminal, No genital ganglion.
like that found in male specimens of P. mugilis.
present.
Systematic position: The specimens fall with-
in the order Palaeacanthocephala Meyer. P.
coorongensis differs from P, mugilis Johnston
& Edmonds, 1947, the latter possessing a
* Department of Zoology, University of Adelaide, Adelaide, S, Aust. 5000.
te GEG ES
mn
Q
Zz
o
2
Q
Ww
e:
‘ Cal gl a et
a O- Imm .-
“ = = aa EE nF mere gt ented
Op TE ee er SE
= ee tay
o
a
~5. Pararhadinorhynchus coorongensis. Fig. 1—Introvert. Fig. 2—Row of nine hooks. Fig. 3.
Figs. 1
Male. Fig. 4.—Posterior of female. Fig. 5.—Egg,
Figs. 6-7. Pararhadinorhynchus mugilis. Fig. 6.—Introvert. Fig, 7.—Egg.
AUSTRALIAN ACANTHOCEPHALA, No, 13 , 21
longer introvert armed with 18 longitudinal
rows of 16-17 hooks per row. It also differs
from Diplosentis amphacanthi Tubangui &
Masilungan, 1937, in which the introvert bears
12 longitudinal rows of 8-9 hooks per row
and in which the Jemnisci are enclosed in a
membranous sac.
is the second acantho-
found in A. forsteri.
aldrichettae | Edmonds
P. coorongensis
cephalan species
Nevechinerhynchus
(1971) is the other.
Pararhadinorhynchus mugilis Johnston & Ed-
monds, 1947: 15.
FIGS. 6~7
The material from which the type descrip-
tion was made contained only two specimens
with everted introverts. Recently a few more
specimens in a fully extended condition he-
came available and consequently the measure-
ments given in 1947 have been rechecked. The
host was the type host, Mugil cephalus Lin-
naeus.
Introvert (Fig. 6) cylindrical. Length of
armed region 0.88-0.94 mm and width 0.16—-
0.25 mm. Sixteen to eightcen, usually 18,
longitudinal rows of 16-17, usually 17, hooks
per row, Unarmed neck, 0,15-0.25 mm long.
Ripe eggs, with polar prolongations of the
middle shell, measure 0.052-0.056 mm x
0.013-0.015 mm.
Type specimens (male and female): Austra-
lian Museum, Sydney (not the South Austra-
lian Museum, as stated by Johnston &
Edmonds 1947).
References
Epmonps, 8S. J. (1972).—Australian Acantho-
cephala No. 13. Three new species. Trans.
R. Soc. S. Aust, 95, 55-60.
Jounston, T. H. & EpMmonps, 8. J. (1947).—
Australian Acanthocephala No. 5. Trans. R.
Sac. S. Aust, 71, 13-19.
Scott, T. D. (1962).—‘“The marine and fresh-
water fishes of South Australia.” (Govern-
ment Printer: Adelaide.)
TUBANGUL, M. A, & MAsILuNGAN, V. A, (1937).—
Diplosentis amphacanthi gen, et sp. nov. an
acanthocephalan parasite in a marine fish.
Philippine Jl. Sci. 42, 183-189.
A NEW GENUS AND SPECIES OF EARTHWORM
(MEGASCOLECIDAE: OLIGOCHAETA) FROM SOUTH AUSTRALIA
BY S. J. EDMONDS*AND B. G. M. JAMIESONT
Summary
EDMONDS, S., J. and JAMIESON, B. G. M., A New Genus and Species of Earthworm
(Megascolecidae: Oligochaeta) from South Australia. Trans. R. Soc. S. Aust., 97 (1), 23-27,
28th February, 1973.
The genus Gemascolex is erected for the new species G. newmani, from South Australia. The genus
is morphologically very similar to Spenceriella Michaelsen, differing primarily in lacking the
extramural calciferous glands of the latter genus. Both genera are assigned to the tribe
Megascolecini. G. newmani is the fourth megascolecid species described from South Australia, but
of those previously described, Megascolex fletcheri Shannon is considered to be a junior synonym
of M. stirlingi (Fletcher), leaving the total number of described South Australian megascolecid
species at three.
A NEW GENUS AND SPECIES OF EARTHWORM (MEGASCOLECIDAE: OLIGO-
CHAETA) FROM SOUTH AUSTRALIA
by S. J. EpMonps* and B. G. M. JAMiEson+
Summary
Epmonps, 5S. J. and Jamieson, B. G. M., A New Genus and Species of Earthworm. (Megas+
colecidue: Oligochaeta) from South Australia. Treas. R. Soe, S. Aust. 97 (1), 23-27,
28th February. 1973.
The genus Gemuscolex is crected for the new species G. newmani, from South Aus-
tralia, The genus is morphologically very similar to Spenceriella Michaelsen, differing primar-
ily in lacking the extramural calciferous glands uf the latter genus. Both genera are
assigned to the tribe Megascolecini. G. riewmani is the fourth megascolecid species described
from South Australia, but of those previously described, Megascolex fletcheri Shannon is
considered to be a junigr synonym of M. stirlinvi (Fletcher), leaving the total number of
described South Australian megascolecid species at three,
Introduction
Three species of Megascolecidae, the only
earthworm family indigenous in Australia,
have previously been named from South Aus-
tralia. They are Megascolex stirlingi (Fletcher,
1888), placed in Perichacta by Fletcher; M,
zieizi Michaelsen, !907a; and M, fletcheri
Shannon, 1920. M. fletcher? Shannon is a
homonym, the binomen having been used by
Michaelsen, 1907a, for u distinct species from
New South Wales. Shannon’s account, though
long, is inadequate in many respects and the
type-specimens are no longer traceable, It
nevertheless conforms sufficiently with the
description of M. stirlingi 10 suggest, though
not unequivocally, that M. flercheri Shannon is
i junior synonym of M. stirlingi. The latter
species has been rediscovered by both authors
in recent collecting in South Australia ‘and
will be described in a future monograph
(Jamieson, in preparation) in which the status
of M, zietzi will also be considered.
The new species erected here has been used
for some years for undergraduate teaching and
for research in the Department of Zoology of
the University of Adelaide. It is referred to
4 new genus which is defined below. Evidence
for inehuding Megascolex stirlingi {Fletcher}
and other species in the new genus will be
deferred to the later publication.
GEMASCOLEX gen. nov.
Terrestrial. Setac numerous (more than 8)
in each segment. A pair of combined male
and prostatic pores on XVIII. Clitellum
annular, anterior to 18/19, Intersegmental
accessory genital markings present. Female
pore unpaired, midventral, in XIV. Spermathe-
cal pores anterior ta IX.
Dorsal blood vessel single. Posterior hearts
latero-ocsophageal, each arising from the sbort
supra-oesophageal vessel and from the dorsal
blood vessel. Latero-oesophageal vessels present
median to the hearts. Subneural vessel absent.
Gizzard large, anterior to septum 6/7. Oeso-
phagus lacking extramural calciferous glands.
Intestine: commencing in XVII, a low ridge
like dorsal typhlosole present; caeca and mus-
cular thickening absent. Excretory system
meronephric. Four pairs of tufted nephridia,
in II-V, their ducts (all?) enteronephric, en-
tering the buccal cavity. Succeeding segments
with astomate, exonephric, micromeronephridia
in lateral bands, Caudally with numerous
enteronephric meronephridia, each with a pre-
* Department of Zoology, University of Adelaide, Adelaide, S. Aust. 5000,
' Department of Zoology, University of Queensland. St. Lucia, Qld. 4067.
24 S. J, EDMONDS and B. G. JAMIESON
septal fiinnel, discharging into the intestine in
each segment. Testes and funnels in X and X1;
testis-sacs absent: seminal vesicles in XL and
XIL
Ovaries and Funnels in XIID, ovisucs absent
Prostates tubuloracemose: lincar, lobulated,
with axial Ittmen rhroughout which reccives
lateral canaliculi; vasa deferentia joining their
muscular ducts. Spermathecue with diverticula,
(Genuscolex, gender male, anagram of Megas-
colex).
Type species: Gemascolex hewmant sp. nov.
Distributions South Australia (see species des-
eription) .
Remarks: The closest morphological affinities
of Gemuascolex lie with two endemic Australian
genera, Spenceriella Michaelsen, 1907b, and
Oreoscolex Jamieson, 1973. The three genera
are the anly members of the indigenous Aus-
tralian Megascolecidae known to possess more
than one pair of stomate nephridia per se2-
ment, and are assienyuble to the tribe Megas-
eolecini Jamieson, 1970. Spencerfella und
Gemascolex unre especially close, as in both,
some wt least of the exudal nephridia are entero-
nephric, opening into the intestine, whereas in
Oreoseolex caudal enteronephry has been de-
duced only very questionably in the type-
species und is unknown in other species, Mul-
tiple nephrostomes and enteronephry have
been demonstrated in Spencerielle for the first
time by Jamicson (in preparation). Oreascelex
tucther differs from Gemascolex and Spen-
ceriella, among other respects, in having only
eight setae per segment.
The chief distinction between Gempscolex
and Spenceriella is the presence of extramural
ciJeiferous glands in Spencerlelfa, Both gencra
differ {rom the type-species of Megescolex in
having stomate nephridia and in their entero-
nephry (Buhl 1946). Only the non-racemose
condition to the prostates would have
separated them from Megascolex in some
former classifications,
Gemascolex newmani sp, nov,
FIG. 1
The following account refers to ihe holo-
type (H)} antl one paratype (P71); variation in
other specimens is discussed subsequently.
Length 180(P1)-234(H) mm, width (mid-
clitellar) 9 mm, number of segments 192(H):
193(P1). Colour in life: dorsal surface brown-
ish purple, ventral surface pale grey; a dark
purplish colour more noticeable at the cx-
tremities; some iridescence present. Cross sec-
tion approximately circular, Prostomium
epilobous 1/3, bisected by 4 longitudinal fur-
TOW, appearing epilanylobaus, owing to longi-
ludinul grooves continuing its lateral limits to
the first intersegmental furrow, but numerous
equally developed parallel grooves present
around the peristomium, Peristomium not
bisected ventrally though in some specimens
a mid ventral groove is more conspicuous than
others. First dorsal pore 4/5. Penchuetine.
setae of cach side more closely spaced laterally
than dorsally and ventrally: Ac slightly wider
than a’, Numbers of setac per segment 32(H)
33(PL) in XU, 26(P1)-30(H) in XX,
31(P1)-32(H) fifteen segments from the
caudal end: a lines straight, z lines irregular;
a ventral and a dorsal break in the setal
eirclet apprectiuble throughowt, Setas a and 6b,
but not ¢, absent in XVI.
INTERSETAL DISTANCES IN
Gemascnlex newmani
~ "standardized =
Te Of citeum ference
tus
tam 6
az oub gy miu aa ab ty ze
Seement NAT
Holutyre 22 08 09 62 23 78 29 32 221
Paratype 1 27 UA 1.0 6.2 28 f2 29 316 RS
Seeneut XX
Holotyne 18 O68 O09 5.6 30 60 20 ju 187
Paratsne t 2.106 13 48 3 7.2 22 42 (8.
Nephropores not externally recognizable.
Clitellum annular, not Eully developed, embrac-
ing XIV-XVIT, but some clitellar modification
present dorsally from 2/3 XII-XVUII
(=5 1/3 segments); intersegments and setae
retained but only # and & conspicuous; dorsal
pores 13/14-17/18 occhided, Mile pores a
pair in XVII, transverse slits in ab(H)} or
b(P1) and almost as wide as ah, each on a blow
transversely Oval papilla, the two papillae out-
lined by a common medianly narrowing field;
the pores 2.8-3.2 mm, 0.08-0.09 circum-
ference apart, Accessory genital markings un-
paired, midventral, transverse, clevated pads,
with lateral limits in ab, in 15/16(H, Pl),
16/16 and 19/20(H), each pad transversed
by a glandular trench corresponding with the
intersegmental furrow but not reavhing to the
ends of the pad, Female pote unpaired, mid-
ventral, in a deep transverse groove at the
anterior border of the setal annulus of XIV_
Spermathecal pores 3 pairs of sunken orifices,
concealed in intersegments 6/7, 7/8 and 8/9,
very shortly lateral of setal lincs a, 2,8=+2,9
mm, 0,09 circumference apart.
Strongest septa 10/11-12/13, very strong.
Dorsal hlood vessel single. continudus onto
the pharynx. Dorsoventral commissural vessels
NEW GENUS AND SPECIES OF EARTHWORM 25
Fig. 1. Gemascolex newmani sp. noy. A, ventral view of right spermatheca of TX (holotype); B
dorsal view of right spermatheca of IX (paratype 2); C, spermathecal pores (holotype); D.
i?
7
wm
>
male genital field (holotype); £, right prostate (paratype 2). Clitellum shaded. All hy camera
lucida. 9, female pore; g.m, accessory genital marking: ¢, male pore; pr.d, prostate duct;
pr.g, glandular portion of prostate; sp.amp, spermathecal ampulla; sp.d, spermathecal duct:
sp,div, spermathecal diyerticulum; sp.p, spermathecal pore. Roman numerals ate segment num-
bers,
in V-XIII; those in X—XIII forming large
latero-oesophageal hearts, each originating
from the supra-ocsophageal vessel but also re-
ceiving, at its junction with the latter, a slender
short connective from the dorsal blood vessel;
these hearts otherwise unbranched. Commis-
surals in V-IX dorso-ventral only, lacking
supra-esophageal connectives, but giving
branches to the posterior septum, gut and lateral
parietes; all commissurals, including the hearts,
yalvular. A pair of Jarge vessels originating on
the parietes in TV passes posteriorly as a pair
of large ventrolateral trunks (latcro-oesophageal
vessels), median to the dorsoventral. commis-
surals, Into TX at the posterior scptum of
which they give branches to the ventral wall
‘of the oesophagus and to the septum, Similar
paired trunks (suboesophageal vessels) present
in XI-XYIJ, closely adherent to the ventral
surface of the oesophagus and lying under its
peritoneum; no continuity demonstrable he-
tween the latero-cesophageal vessels in IX an-
™ S. 1, ERBMONDS und B. G. JAMIESON
teriorly and the suboesophageal vessels in XI
posteriorly, both pairs of vessels give a pair of
vessels to the nesuphageal plexus in each seg-
mene. Submeural vessel absent, Gizzard moder-
ately lurge, fusiform, and firrnty muscular in
Vii septum 5/6 adherent to its anterior end:
its posterior end projecting a little behind inter-
segment 7/8; the ocsophagus much narrower
in V, short and narrow in VIL dilated and
vascularized with low intermal rugac, but ne
calciferous glands, in VUI-XY, narraw and
short in XVI Intestinal origin XVIL; a low
ridgelike dorsul typhlosole commencing in
XXIF, caeca and muscular thickening absent.
Nephridia: meronephric: large paired tufts,
with very many spiral loops, in I. IL. TV and
V increasing in size pasteriad and very
large in V. Those in V and IV sending antero-
medianly thick sheaves of numerous ducts,
which loosely aggregate ay 4 composite duct
common to both pairs of tufts, the ducts pass-
ing forward (to join the lateral wall of the
buecal cavity in front of the brain, Those in
Il and I sending slender composite ducts to
the lateral walls of the buccal cavity immed-
iately behind the muuth, Nephridia in succecd-
ing segments astomate. exonephric, micro-
meronephridia: very dense transverse bands of
spital tubules, which laterally may be ¢on-
sidered to form tufts, anterior in WI, their
numerous discrete or partly aggregated ducts
Hischarzing exonephrically at the anterior limit
of the segment; some exonephric nephridia
present! pasteriorly in the segment; VIT-X each
with an anterior and a. posterior parietal band
of numerous nephridia; XI, XU and XT with
similar but rather sparse bands; XIV with
sparse anterior and dense posterior bands; XV
with Uenser unierior und sparse posterior
bands; XVI (the last oesephages! segment)
and succeeding intestinal segmests with only
anterior hands; hands dense in XIV-XVI1L.
Caudally, commencing at segment 120 (in
holotype with [92 segments), with numerous
enlarged onephridia (almost megamerone-
phridia) in each segment on the anterior wall
of the segment near to ard encireling the in-
testine, the nephridia about five deep and cach
with 4 long-slalkecl conspicuous. preseptal
funnel, the nephridial ducts in cach segment
Tunning medially as a common duct on each
side to enter the wall of the intestine anteriorly
in the segment on each side of the dorsal blood
yessel, though some individual nephridial ducts
reich the wall of the intestine independently in
the vicinity of their nephridial hodies, Concen-
tric with and external io the enlarged, entero-
nephric nephridia are parietal astomate,
apparently ¢xonephric, micromeroncphridia in
dense transverse bands. The number of cn-
larged nephridia decreases, and that of the
micromeronephridia greatly increnses, m the
last twenty or so segments.
Holandric; sperm funnels iridescent in X
and XI; Lestis-sacs absent; seminal vesacles
large lobulated sacs im XI and XII; similar.
smaller, structures on the anterior septum of
X ate of unknown function but cannot be
seminal vesicles. Ovaries (webs of large
oocytes) and funnels in XIII; vvisics nbsent,
Prostates limited to X VINE, flattened, Jobulateel
S-shaped glands with short slightly tortuous
muscular ducts joined near their ental ends by
the vasa deferentia. Penial setac absent.
Spermathecae 3 pairs, in VU, VII and IX.
diveruculun (inseminated) single, subspherical.
internally multiloculate, with a shert stalk (H,
Ph).
Field Variation: In 16. clitellate type-speci-
mens. including the holotype, a lransvetse
median genital marking is present in [5/16
and 16/17 in 9 specimens, and in 19/20) in
fO specimens, Seven of these specimens have
the Full complement. of markings, in 15/14.
16/17 and 19/20; 2 specimens have the geni-
tal marking m 15/16 only; 2 have them in
16/17 and 19/20 only: 1 has a marking in
19/20 only; und 4 have no murkings,
Murerial Exentined: Cudlee Creck, 34° 50°S_.
138°49°E,, from helow apple orchard of Mr.
G. Newman, Edmonds, August 1971—H,
PI-9; N, Maier, 21L.viti1972—PlI-s3.
Hahndorf, 35°02'S., 138°48'E.. G. Perersan,
25.viii 1972—P 70-12.
H, P2-4. deposited in the Australian
Museum. Sydney; PL. 5, British Muscum
(Natural Histury); P6. 7, South Australion
Mailscuum; P8—-12 famieson collection,
Remarky: Material used in undergraduate
stadies and not retained cannot be designated
type-material but. nevertheless, variation in it
may be noted. In 50 specimens the length was
200-271) mim; the width was 7-9 min generally
but 8-11 mm in the region of crop and gizzard.
The clitelum embraced XIV-XVII hui some-
times included part of MII and of XVIIL
Numbers of spetmathecal pores and location
of the male pores and of the female pore were
constant. Accessory genital markings, when
present, were usually at 15/16, 16/17 and
19/20, The iitestine usutaily began. iy XVI,
a condition also noted in some nf the type-
NEW GENUS AND SPECIES OF EARTHWORM 37
specimens. The flattened S-shaped form of the
prostales was constant, The supra-oesophageal
vessel occupied VII-X1V, ramifying on the
oesophagus at each end, In one hundred speci-
mens the number of segments was 155-198,
The course of the suboesophageal blood
vessels has not been unequivocally determined
and, with that of the latcro-parietals, requires
further examination. The structures resembling
seminal vesicles anteriorly in X do not show
spermato-genests in serial sections, whereas the
seminal vesicles in XI and XII do.
Gemascolex newmani is distinguished from
the equally large Megascolex stirlingi, so fai
as it is described by Fletcher, 1888, in (a)
location of the spermathecal and male pores
in ab lines whereas in M. stirlingi they are
markedly more lateral, in setal lines 6 and be-
tween setal lines 3 and 4 respectively; (b) the
unpaired instead of paired accessory genital
markings and their forward extension to inter-
segment 15/16; (c) the S-shaped, not straight,
prostates and in other respects.
Noteworthy differences from MM. zierzi, as
described by Michaelsen, 1907a, are (a) the
more median location of spermathecal and
male pores, (b) location of the male pores well
lateral of paired accessory genital markings in
M zietzi, (¢) presence of further paired mark-
ings on the anterior border of the male poro-
phores in the latter species; (d) restriction of
the prostates to XVIII and their S-shaped
form: (c) the subspherical sessile spermathe-
cal diverticulum of newsani contrasted with
the very Jong tortuous tubular diverticulum of
M. zietzi.
Acknowledgements
The authors thank Mr, G, Newman for per-
mitting collection of the new species from his
property and Mr. G, Peterson and N. Maier
for procuring additional material. This work
was carried out during tenure of ARGC and
URG grants,
References
Bani, K. Ny (1946).—Studies on the structure,
development, and physiology of the nephridia
of Oligochaeta. Part VII. The enferonephric
type of nephridial system in carthworms be-
longing to three species of Megusvolex
Templeton and three. species of Travoscolides
Gates (Meguscolides McCoy), Q. JI microse:
Sei. 87, 45-60,
FLeTcHer, J. J. (1888)—Notes on Australian
earthworms, Part II, Proc. Linn. Soc, NSW.
2, 375-402.
Jamirson. B. G. M. (1970).—A review of the
megascolecoid earthworm genera (Oliza-
chasta) of Australia, Part 1—Reclassification
and checklist of the megascolecoid genera of
the world. Prov. R. Soe. Od B2(6), 75-86.
JAMizson. B. G. M,. (1973).—Earthworms
(Megascolecidac: Oligochueta) from Mount
Kosciusko, Australia. Rec. Ast. Mus. (in
press.) .
MICHAELSEN, W. (1907a).—-Oligochaeten von Aus-
tralien. Abh. Geb. Naturw., Tamburg 19(1),
1-25.
MICHAELSEN, W. (19076) —Oligochaeta in: Die
Fauna Sidwest-Australieny 1(2), 177-232.
SHANNON, J. H. (1920).—On the structure of 2
new species of carthworm from South Aus-
tralia, Megascolex fletcheri. Proc, R. Soc.
Viet. 32, 301-313, Pl. XXVII-XXXI
AERIAL DISPERSAL OF ADULT CARDIASPINA DENSITEXTA
(HOMOPTERA:PSYLLIDAE) IN SOUTH AUSTRALIA
BY T. C. R. WHITE*
Summary
WHITE, T. C. R., 1973. Aerial Dispersal of Adult Cardiaspina densitexta (Homoptera:Psyllidae) in
South Australia. Trans. R. Soc. S. Aust. 97 (1), 29-3 1, 28th February, 1973.
Living adult C. densitexta were captured at 152 m and 305 m above the ground in the vicinity of
Keith, South Australia during the spring. This finding supports the hypothesis that the adults of the
spring generation generally function as the effective dispersive stage in the life cycle of this insect.
AWRIAL DISPERSAL OF ADULT CARDIASPINA DENSITEXTA (HOMOP-
TERA: PSYLLIDAE) IN SOUTH AUSTRALIA
by T. C. R. WuiTeE*
Summary
Wurrr, T. C. RK. 1973, Aerial Dispersal of Adult Cardiaspina densitexta (Homaptera:
Psyllidac) in South Australia. Trans. R. Sac. §. Aust. 97 (1), 29-31, 28th Febrmary_ 1973.
Living adult C. densitexta were captured at 152 m and 305 m ahove the ground in the
vicinity of Keith, South Australia during the spring. This finding supports the hypothesis that
the adults of the spring generation generally function as the effective dispersive stage in the
life cycle of this insect.
Introduction
Clark (1962) concluded as a result of his
obseryations and experiments with adults of
Cardiaspina albitextura Taylor that emerged
in the autumn, that they tend not to disperse
far from their point of origin, and that there-
fore this is “ . not a strongly dispersing
insect.”
Studies of the closely related Cardiaspina
dénsitexta Taylor (White 1970a) suggested
that although this seemed to he true of the
generation of adult C. densitexta that emerged
in the summer and autumn, it did not seem to
be true of the generation that emerged in the
spring. During summer and autumn, trees that
were isolated from the established infestations
were rarely colonized, Also at this time of the
year. adulis seemed to show great “reluctance”
to leave the foliage on which they had devel-
oped, On the contrary, during spring it was
commonplace for isolated trees to be colonized
quickly. Also direct observation of the be-
haviour of adults at this season of the year
showed that they had a strong tendency to
fly up and away from the foilage on which
they had lived as nymphs.
These observations led to the hypothesis
that C. densifexta exhibits two types of seasonal
behaviour: (1) non-dispersive, serving to con-
Sentrate summer and autumn adults—and sub-
sequent egg-laying—on favourable foilage; (2)
dispersive, serving to distribute the adults
emerging in ihe spring away from depleted
foilage ind increasing the chance of some of
them finding fresh foilage.
This would suggest that the adults thai
emerged in the spring were the effective dis-
persive stage in the life cycle of C. denvirexta,
most probably in the manner proposed by
Lewis & Taylor (1965). Their analysis of
many aerial samples demonstrated that “high
altitude and long distance migration is very
highly correlated with flight by day and small
size.”
Such an hypothesis presupposes that adults
of C. densitexta which emerged in the spring
were present high in the air where they would
be widely and randomly dispersed by air cur-
rents and winds. In 1964 an attempt to demon-
strate this was unsuccessful. By the time a
suitable technique had been developed, num-
bers of C. densitexia had fallen to very low
levels, reducing to near zero the probability of
catching the few individuals that might have
been present in the aerial plankton,
This paper Treports a second attempt to
catch airborne adult C. densitexta at a time
when they were abundant.
Materials and Methods
Nets mounted on a steel ring 38 cm in
diameter were towed from the wing of a
Cessna 172 aeroplane flying at 1352 and 305 m
above the ground. Details of the construction
* Department of Zoology, University of Adelaide, Adelaide, S. Aust. 5000.
Present Address; School of Natural Resources, University of the South Pacific, Sova, Fiji,
30 TCR,
of the nets and the cing, and of the method of
Operaling, were described previously (White
1970b).
All samples were of 15 minutes duration.
Euch net was released when the aeroplane had
levelled off at the required height and was
cruising at 121-131 km/h. Height and speed
were maintained as accurately as possible for
the duration of each sample.
All nets were kept sealed in plastic bags
until immediately before being used, At the
completion of each sample a knot wus tied in
the net before it was detached from the ring
and resealed in w plasti¢ bag. All bags were
returned to the laboratory before being opened
and the nets immersed in 70% alcoho! before
the knots were untied.
All collections were made over a relatively
amall area between Keith and Willalooka jin
the southeast of South Australia (White 1970b,
fig. 2), At the time the host plants (Eucalyp-
ius fascieulosian Fv.M.) in this locality were
carrying w moderate to high population of C.
densitexta, wind ahout 50%. of the adults had
emerged.
Results
Forty-one samples {Table 1) were collected
during, five days, thirty at 152 m and eleven at
305 m, Twenty cight contained adult C. den-
xitexia, the catch varying from one to nine
per sample at 152 m, and one to five per
sample at 305 m, A total of 60 adults were
caught. there being approximately equal num-
hers of males and females, Several were ob-
served crawling around in the nets after these
had been placed in ptastic hags.
TABLE |
The number of adult C_ densitexta captured
No. of Aln-
Date Samples — tude a 9 + Fatal
Lexi7t LL(7j* [52m 5 5 1 oul
2.x1.71 77) 1S2m fi 8 2 16
3x17] §(3) 132m 2 2 1 3
VL .xi.7t 4(3) [52m 4 5 2. al
4(4) wim — 2 ] 3a
12.xi.71 73) 305 m 5 of 3. 614
Total 41(28) _— 22 28 #10 60
* Number of samples containg C. denstrexta in
parenthesis.
+ Damuged individuals lacking an abdomen,
A net with a diameter of 38 cm, pulled
through the air at a mean speed of 126 km/h,
would sift 221 m* of air per minute—3319 m!
in a 15 minute sample. But because of the
WHITE.
“cushion” of air formed in Front of the net. at ix
improbable that this volume of air would have
passed through the net, Much of it would have
been deflected around the sides of the net, No
attempt was made to assess the “effective”
diameter of the nets—it. may well have been
no more than 2.5 om to 5 ¢m but if a conser-
vative 7.5 em miurgin is deducted to whow for
deflection an clfevtive diameter of 23 ¢m re-
mains. Then 1188 m® of air would have been
sampled in 15 minutes.
A 152 m high block of air ubove a hectare
of jand contains 1,520,000 m* of air, at 305
m, 3,050,000 m*, Tf the adult psyllids were
distributed wt random through this air, the
sampling indicated densities of from about
1,300 to 11,500 per hectare to a height of 152
m and trom 2,500 to 12,800 per hectare to
305 m.
Discussion and Conclusion
The number of unknowns makes any
uttampt to quantify the density of adult psyllids
in the aerial plankton unrealistic. Quite apart
trom the problem of the effective size of the
sampling unit, it is likely that the distribution
of ait-borne psvilids was far from random.
The time of day, ambient temperature, wind
speed and the extent and duration of the warm
au “thermals” rising from the ground would
all have contributed to a patehy and variable
distribution, both vertically and horizontally,
Almost certainly this sampling underesti-
mated the number of these insects in the air
over any area of land, But equally certainly it
demonstrated that there were very many of
them carried to considerable heights above
the trees on which they emerged; sufficient to
ensure that they would have been scattered
over many square km of land, As the air
cooled each evening and they returned to earth.
relutively few of them would have been for-
junate enough to Jand on a suitable host plant.
But the few that did would have survived, and,
with their huge capacity for increase. soon
utilized the available food.
The evidence from this sampling. combined
with that previously reported {White 1970a),
demonsteated that the adults of C. densitexta
that emerge in the spring are the means by
which this species of psyllid i widely and
effectively dispersed.
Other records of adult psyllids captured high
in the air (Glick 1961; Freeman 1945, Hardy
AERIAL DISPERSAL OF CARDIASPINA DENSITEXTA 31
& Milne 1938) and as far as 298 km from the
nearest land (Yoshimoto & Gressitt 1963;
Harrell & Yoshimoto 1964) suggest that this
method of dispersal is common within the
Psyllidae.
Acknowledgements
A grant from the Endowment and Scientific
Research Fund of the Royal Society of South
Australia made it possible for this sampling to
be carried out.
I am grateful to Gordon Burnard for
arranging the hire of the aeroplane from the
Pinnaroo Aero Club and for his and Barry
Browne’s patient and skillful piloting. To Joan
Burnard my thanks for her hospitality and for
allowing her kitchen to be converted into a
temporary laboratory,
References
CLaRK, L. R. (1962).—The general biology of
Cardiaspina_ albitextura (Psyllidae) and its
abundance in relation to weather and para-
sitism. Aust. J. Zool. 10, 537-86.
FREEMAN, J, A. (1945).—Studies in the distribu-
tion of insects by aerial currents, The insect
populations of the air from ground level to
300 ft. J. Anim. Ecol, 14, 128-54.
Guicx, P. A. (1961).—Airborne movement of the
Pink Bollworm and other arthropods. Tech,
Bull. U.S. Dept, Agric. 1255, 1-20.
Harpy, A. C. & Ming, P. S. (1938).—Studies in
ihe distribution of insects by aerial currents.
Experiments in aerial tow-netting from kites.
J, Anim. Ecol. 7, 199-229,
Harrei, J. C. & Yosuimoto, C. M. (1964).—
Trapping of airborne insects on ships on the
Pacific, part 5. Pacific Insects 6, 274-282.
Lewis, T. & Taytor, L. R. (1965).—Diurnal
periodicity of flight by insects. Trans. R.
ent. Soc, Lond, 116, 393-479.
Waite, T. C. R. (1970a).—Some aspects of the
life history, host selection, dispersal and
oviposition of adult Cardiaspina densitexta
Comaninrs = Feyiligar). Aust J. Zool. 18,
105-17,
Waite, T. C. R. (1970b).—Airborne arthropods
collected in South Australia with a drogue-
net towed by a light aircraft. Pacific Insects
12, 251-259.
Yosutmoto, C. M. & Gressirr, J. L, (1963).—
Trapping airborne insects in the Pacific-
Antarctic area, 2. Pacific Insects 5, 873-883.
LIFE HISTORY, LARVAL MORPHOLOGY AND RELATIONSHIPS OF
AUSTRALIAN LEPTODACTYLID FROGS
BY G. F. WATSON AND A. A. MARTIN®*
Summary
WATSON, G. F. and MARTIN, A. A., 1973. Life History, Larval Morphology and Relationships of
Australian Leptodactylid Frogs. Trans. R. Soc. S. Aust., 97 (1), 33-45, 28th February, 1973.
Disagreement exists regarding the phylogeny, relationships and classification of Australian
leptodactylid frogs. Analysis of their life history patterns indicates that one of the present two
subfamilies, the Myobatrachinae, is a close-knit natural group, whereas the other, the Cycloraninae,
is more heterogeneous. In particular, the genus Cyclorana does not conform with the cycloranines,
and in terms of life history has strong affinities with the Hylidae. No close relationship between the
Myobatrachinae and the Cycloraninae is evident from life history data.
LIFE HISTORY, LARVAL MORPHOLOGY AND RELATIONSHIPS OF
AUSTRALIAN LEPTODACTYLID FROGS
by G. F. WatTson* and A. A, MartTIn*
Summary
Wwarsan, G, PF, and Martin, A. A., 1973. Life History, Larval Morphology and Relationships
of Australian Leptodactylid Frogs. Trans, R. Soc. S. Aust., 97 (1), 33-45, 28th February,
1973,
Disagreement exists regarding the phylogeny, relationships and classification of Australian
leptodactylid frogs. Analysis of their life history patterns indicates that one of the present
two subfamilies, the Myobatrachinae, is a close-knit natural group, whereas the other, the
Cycloraninae, is more heterogeneous. In particular, the genus Cyclorana does not conform
with the cycloranines, and in terms of life history has strong affinities with the Hylidae. No
close relationship between the Myobatrachinae and the Cycloraninae is evident from life
history data.
Intreduction
In the first substantial monographic study of
Australian anurans, Parker (1940) divided the
Australian representatives of the family Lepto-
dactylidae into two subfamilies: Cycloraninae
and Myobatrachinae. The major characters
used to define the twa groups were the struc-
ture of the tongue, hyoid apparatus, larynx
and thigh musculature, Martin (19672) noted
that biological characteristics, particularly life
history, were broadly consistent with Parker’s
division, Lynch (1971), using a complex of
morphological, osteological and ecological
characters, substantiated Parker's taxonomic
interpretation; he further divided Puarker’s
Cycloraninae into two tribes, Cycloranini and
Limnodynastini. Tyler (1972a) investigated
the superficial mandibular musculature and
vocal suc struciure of Australian leptodacty-
lids, and came to conclusions similar to those
of Parker and Lynch, with one important ex-
ception, Tyler found that Cyclorana Stein-
dachner did not conform with either the
Cycloraninae or the Myobatrachinae; and he
questioned its familial disposition, noting that
it shared some characters with the amily
Hylidae.
The generic classification of Australian
Jeptodactylids has undergone considerable
modification since Parker's work. Heleioporus,
as recognized by Parker, was divided into two
genera, [eleioporus Gray and Neobarrachur
Peters. by Main (1957a); and the new genera
Kyarranus Moore, 1958 and Taudactylus
Stranghan & Lee, 1966 have been erected.
Tyler (1972b) removed Crinia darlingtoni to
# new genus, Assa. Blake (in press), using a
polythetic mumerical approach; finds that
Crinia is divisible into three gencta, and group-
ings corresponding to these genera are used
here, They are referred to as the Crinia has-
well’ group (including C. haswelli and C.
georgiana); the Crinia laevis group (including
C, Jaevis, C. leai, C. lutea, C. rosea and C.
victoriana); and the Crinia signifera group (in-
eluding all other species of Crinia). Blake (in
press) also finds that Mezacrinia Parker does
not warrant separation from Pseudophryne
Fitzinger and he intends to synonymise these
two genera; hence they are not treated separ-
ately here. The current composition of the
Australian leptodactylids is shown in Table 1.
The present contribution summarizes the
available data on the life histories and larval
morphology of Australian leptodactylids. Such
information is useful from two points of view,
First, life history stages provide morphological
characters independent of those exhibited by
+ Department of Zoology, University of Melbourme, Parkville. Vic. 3052.
a4 G. F. WATSON and A, A, MARTIN
adults, and can therefore be uscd to test
phylogenetic relationships based on studics of
adults (such gs the suggested hylid affinitics
of Cyclorana), provided that care is taken
to recognize convergence (Inger 1958; Griffiths
1963}. Second, life history is an indicator of
geferal adaptive ecology, and generic defini-
tion is currently based on ecological as well as
morphological characters (Mayr 1969). Inger
(1958) has demonstrated the utility of life
history data in anuran classification at the im-
frafamilial levels, and this approach was used
by Martin & Watson (1971) in an analysis. of
the family Hylidae. Although Lynch (1971)
took life histories into account in his analysis
of the leptodactylids, mast of his dats on Aus-
trilian forms as derived from the literature,
and is often inadequate or erroneous.
TABLE |
fnfrafamiltal classifications of Astralian
leprodavtytids,
Adelotun
Kyarramis q
Lechriodus Linney
Lim nodynuntcs: fork 197) Subsamii
" ft
Philovin Cycloraninae
Tribe |
; F Yas
Meleigporus | Bates
Subfamils
Cyeloruninac
(Tyler 19729)
Mixophyer Tribe
Neobatachut Cycloranini
Notaden | cad 171)
Cyclorans
Crinia laevis group
Crinia signitera group
Glavertia
Myobarrachus
Peeudophryne
Taudactylus
Uperoleia
Subfamily
Myobatruvbinar
(Varker 1940; Lyaet 197t,
Asia
Crinia haswelh sroup |
f Teler 19720; Make in prose)
Material and Methods
Material representing all but four of the 17
venera of Australian leptodactylids has. been
examined, The four gencra not studied are
the cycloranine Noteden Gunther ahd the
myobatrachines Awsa Tyler, (lanertia
Loveridge and Myehatrachuy Schlegel. For
these genera data have been drawn from the
literature. One or more species of each of the
other genera have been examined; all obser
vations mot supported by 4 reference are
original. In some cases (e.2, Limnoadynastes,
the Cvrinia signifera group) enough species
have been stucbed to be fairly confident that
the limits of intrageneric vartalion in life his-
tory have been detected; in others (¢,g. Tanda
tylus), only one species hus been examined
and the present account may therefore not be
characteristic of the genus.
Identity ul life history stages was established!
hy rearing eges of known parentage, or by rais-
ing to metamorphosis a portion of each tuilpele
sample, The only exception is Taudacrylus,
where identification of the larvae is based on
the fact that they were collected at the type
locality of 7. diurnus, and are distinct fram
the larvae of any other anuran known to in-
habit the area.
Although data on numerous characters were
assembled, five major feutures of the life
history showed consistent variation and were
employed in our analysis, These features, inost
of which are illustrated in Martin (1965), are:
(1) Type of egg mass: whether foamy or
nat.
(2) Larval development: whether terres-
trial or aquatic.
(3) External gills: whether present or ab-
sent in embryonic development,
Number of rows of teeth in the upper
lubium of the larval mouth: whether
nonc, two, or more than twa.
Disposition of labial papillae in the
larval mouth; whether completely sur-
rounding the mouth disc (na gaps), or
with an anterior gap, or with both .an-
terior and posterior zaps.
One additional character commonly em-
ployed in larval descriptions—whether the
anus is median or dextral—is included for the
sake of complcteness, but in many groups 7
is too variable (even within genera) to be use-
ful in the analysis of affinity (Lee 1967; Lynch
1971). The larval morphology of each genus
is Wustrated by drawings of the indpoles of
one or more species in lateral view, and of the
larval mouth dises, Larvae between stages 3f)
und 38 of Giosner (1960) were used for illu-
stration. Where material was avilable For
several species in a genus, the specics selected
for illustration and description is generally
one which has not been considered in our pre-
vious publications (e.g. Martin 1965, 19674),
The description relers to the species illustrated,
and variations in other species are noted.
Drawings were mude with the aid of a stereo-
scopic microscope, using photographs, a
cameta lucida, or an ocular micrometer and
squared paper.
(5)
Survey of Life Histories
Adelotus Ogilby
Species examined: A. Arevis, Trani Mt_ Nebo.
Old.
The egg mass is foamy und is deposited In
standing or flowing water, development 4s
AUSTRALIAN LEPTODACTYLID FROGS
Sie me,
Cree
ea epE ITS os,
C Pepa a ee
ey my
S97 > pcean,
SPER
Fig. 1, Left lateral view of larvae of: A, Adelotus brevis; B, Lechriodus fletcheri; C, Kyarrunus
sSphagnicolus;
45mm,
aquatic. The eggs lack pigment and have a
diameter of 1,7 mm (Martin 1967a). There
are no external gills. The anus is dextral, and
the larva has an unspecialized body form (Fig.
1A). There are three upper and. three lower
rows of labial teeth (not two upper and three
lower as stated by Lynch 1971), and a gap in
the labial papillae along the anterior margin
of the mouth disc (Fig. 2C),
Adelotuy is a monotypic genus.
Kyarranus Moore
Species examined: K- sphagnicolus, from Point
Lookout, N.S.W.
The foamy egg mass is placed out of water,
in damp sphagnum moss, and the larvae do not
feed, though they may become free-swimming
(J. M. de Bavay, pers. comm.). Small external
gills are present, The larva has a relatively long
tail and broad fin; the anus is median (Fig.
1C). The mouth parts are reduced, with well-
developed jaws but no labial teeth; the papillary
border is broken anteriorly (Fig. 2D).
The life history of K. loveridgei is very
similar (Moore 1961).
BD, Limneodynastes interloris; E, Philoria frosti, In each case the bar represents
Lechriodus Boulenger
Species examined: L. flercher?, ftom Cunning-
ham's Gap, Qld.
Development is aquatic, often in highly
ephemeral situations. The egg mass is foamy
and the ovidiameter is about 1.7 mm (Martin
1967a). Long, filamentous external gills are
present. The larvae (Fig, 1B) are carnivorous
and development is rapid (Moore 1961), The
anus is median. The mouth disc (Fig. 2A)
has large jaws, and six upper and three lower
rows of labial teeth. Labial papillae are absent
from the anterior margin of the mouth disc.
Lechriodus is incorrectly described by Lynch
(1971) as having only two upper labial tooth
rows,
L. flercheri is the only representative of the
genus in Australia, but there are several species
in New Guinea (Parker 1940).
Limnodynastes Fitzinger
Species examined: L. dumerili, L. fletcheri, L
interioris, L. peroni, L. salmini, L. tas-
maniensis, L. lerraereginae.
36 G. F. WATSON and A. A. MARTIN
i
. Panis
we
~™
psy ree
‘ust tle eV,
we Er alee ge Oh
veel apnea aytyner en
ee eee
a eg
com a UA
~ ah
mM
rt
‘aye
" ven eH Ye nny r
to a
x 1 ee Nara wh a
we bingy SILAS
VERA VEY ITER S
PIAVIL ERAN
—oe
My
yi
ca
%
MT
Nanya NTE
e
Fig. 2. Laryal mouth discs of: 4, Lechriodus flétcheri: B, Limnodynastes interioris; C, Adelotus brevis;
D, Kyarranus sphagnicelus; FE, Philoria frasti. In each case the bar represents 1 mm,
Species deseribed: L. inrerioris, from Boree
Creek, N.S.W.
The egg mass is large and frothy, and is
deposited in water among vegetation, under
logs or rocks, or in water-filled burrows in
stream banks (Martin 1967a); development is
aquatic. The ovidiameter is about 1,7 mm.
Externat gills are small and unbranched. ‘The
larva (Fig. 1D) has a generalized body form;
the anus is median, There are six upper and
three lower rows of labial teeth, and Jabial
papillae suzround the mouth disc except for
the anterior margin (Fig, 2B).
This life history pattern seems fairly con-
stant throughout the genus. In southern Vic-
torign and Tasmanian L. peroni, the eggs are
unpigmented (Littlejohn 1963a). Egg counts
range from 1,100 in ZL. tasmaniensis to 3,900
in L. dumerifi (Martin 1967a), All species
have at least 4, und usually S—6, rows of teeth
in the upper labium.
Philoria Spencer
Species cxamined: P, frosti, from Mt, Baw
Baw, Vic.
AUSTRALIAN LEPTOGDACTYLID FROGS 37
ET gl,
fig SH
Fig. 3. Left Jateral view of larvae of: A, Heleioporus australiacus; B, Cyclorana cultripes; C, Mixe-
phyes balhus; D, Neobatrachus pictus, Tn each case the bar represents 5 mm.
The frothy egg mass is placed in damp
sphagnum near water; the eggs are unpig-
mented and have a diameter of 3.9 mm (Liftle-
john 19636). Small external gills are present.
The Jarvae may be free-swimming but
apparently do not feed, The anus is median
and the tail fin large in proportion to the body
(Fig. 1B). The mouth has well-developed
jaws, and papillae on the Jateral and posterior
margins of the disc, but labial tecth are absent
(Fig. 2B).
Philoria is a monotypic genus.
Heleiaporus Gray
Species examined: H. awstraliacus, ftom 12
km 'S. of Walhalla, Vie.
The eggs are unpigmented and measure 2.6
mm in diameter; egg counts of four masses
ranged from 775—1,239 eggs. The egg mass
is foamy and is deposited in standing or flowing
water concealed in vegetation or in burrows;
development is aquatic. The external gills are
prominent. The Jarvae (Fig. 3A) are un-
specialized, with a median anus. The mouth
disc has six upper and three lower rows of
{ahial teeth, and an anterior gap in the papil-
lary border (Fig. 4A).
lee (1967) has described the lite histories
of the five Western Australian species of
Heleivporus. In these, the cggs (mean ovi-
diameters 2.6—3,8 mm; mean egg counts 160—
480 eggs) are laid in dry burrows which are
later flooded, and the larvae undergo aquatic
development. The anus muy be median of
dextral. and there are 5-6 rows of teeth in the
upper Jabium-
Mixophyes Gunther
Species examined; M. halbus, M. fasciolatus.
Species described: AM]. balbus, from Point Look-
out, N.S.W.
The cggs (ovidiameter about 2.8 mm) ure
pigmented and are laid in clusters on rocks or
gtavel near the edge of flowing streams, Each
egg has a distinct separate capsule, and the
mass 3s not frothy, External gills are present.
Development is aquatic, and the larva is a
large and powerful lotic form (Fig, 3C). The
anus is dextral. The mouth disc has a com-
plete papillary border, and six upper and three
lower tows of jabial teeth, There are also 5-6
lateral rows on each side near the angle of the
jaw (Fig. 4B). The hind limbs develop in a
membrgnous sac and are not visible until late
in development.
The life histories of Mf. balhus and M-
fasciolatus appear to be essentially identical
(Martin 19674). Details of life history in the
other two members of the genus are not
recorded,
Neobatrachnus Peters
Species examined; N. centralis, N. pictus.
Species described: N. pictus, from Savernake,
N.S.W.
The eggs are pigmented and about 2.2. mm
in diameter. They are Jaid in strings of jelly
wound among submerged vegetation in stand-
ing water, and development is aquatic. Small
external gills are present. The larvae are active
swimmers with relatively plump bodies.and short
§ G, F. WATSON
—_——> 7
SSP mht ee
Teac ee Cr <
nm al Arnaut sen aie
POLLEN
sry naan nen oo
Ae
and A, A. MARTIN
Seti,
n Uh Sener a,
yltss f
iL
atl
ek
RA Usyas 14-4
Peery yan?
a,
acl ; re
ST
4 ee
a gabtirae ing
Fig. 4. Larval mouth discs of: A, Heleioporus ausiraliucus; B, Mixophyes halbus; C, Neobatrachts
pictus; D, Cyclorana coltripes. In each case the bat represents 1 mm.
jails (Fig. 3D). The anus is median or slightly
displaced to the right. Papillae are absent from
the anterior margin of the mouth dise, and
there ate three rows of teeth in each labium
(Fig. 4C). The jaws are very robust, presum-
ably reflecting the fact that the larvae feed by
ingesting large fragments of plant and insect
material (M. J. Tyler, pers. comm.).
In castern populations of N. centralixs the
eygs huve discrete capsules and are laid sep-
arately or in loosely adherent clumps. The
three endemic Western Australian species lay
their eggs in Jong strings (Main 1965, 1968).
Notaden Gunther
Species deseribed: N. richollsi, from Munka-
yarra, W.Aust.
This account is taken from SJater & Main
(1963). The eggs are 1,3 mm in diameter and
pigmented; they are laid in temporary pools
and development is aquatic. The form of the
egy mass, and whether or not external gills
develop, are not recorded. The anus is median.
There ate three upper and three lower rows
of labial teeth, and papillae extend around the
sides and back of the mouth disc.
Vhere is no information on record concern-
ing the life history of the other members of
this genus, N. hennetti and N. melanoscaphus.
Cyclorana Steindachner
Species examined: C. australis; C, cultripes, C,
platycephalis.
Species described: C,
Creek, N.T.
Development is aquatic, "The eggs are small
and pigmented and are Jaid in clusters, without
distinct separate capsules, in water (Main
3965), Embryonic development is not re-
corded, The larva (Fig. 3B) has a distinctive
acuminate tail tip; the anus is dextral, though
often only slightly displaced from the midline.
There are two upper and three lower tows of
labial teeth, and papillae occur along the
lateral and posterior margins of the mouth
disc (Fig. 4D).
The eggs and larvae of C, platyeephalus,
and the larvae of C. australis, are. similar to
those of C, cultripes. No data are availible for
other species in the genus.
culrripes, from Pine
AUSTRALIAN LEPTODACTYLID FROGS wy
a
eer
Fig. 5. Left lateral view of larvae of: 4, Crinia kaswelli; 2, Psendophryne corrobaree: C, Crinia leeviy:
DBD, Crinia rasea. In each case the bar represents 5 mm.
Assa Tyler
Species described: A. darlingtoni, ‘rom the
Macpherson Range. Qld.
The eggs are unpigmented and average 2.5
mm in diametet. Oviposition and embryonic
development are not recorded. The larvae de-
Velop in a brood pouch of the male, but details
of their morphology have nor been described
(Straughan & Main 1966).
The genus A'vsa is monotypic.
Crinia Tschudi
‘CRINIA HASWELLI group
Species examined: C. Aaswelfi, from 7 km W.
of Orbost, Vic,
The eggs measure about 2 mm in diameter
uid are pigmented, with distinct individual
capsules. They are laid in water and develop-
ment js aquatic. External gills are absent. The
larva (Fig. 5A) is a specialized nektonic form
with high fins. The aous is dextral. The tadpole
appeurs to feed largely on plankton, The
mouth has two upper and two lower rows of
labial teeth, and a single row of papillae bor-
dering its lateral and posterior margins (Fig.
6A).
The Western Australian C. georgiana, the
only other member of this group, has 4
markedly different pattern of development
(Main 1957b, 1965). The eggs ate latd in
permanent streams and soaks. Larvae are of
the lotic type, being flattened, and with long,
slender tails (see Main 1957b, Fig. 2a).
There are three rows of teeth in egch labium,
and the papillary border has both anterior and
posterior gaps.
CRINIA LAEVIS group
Species examined: C. laevis, C. resea, C. vie-
7oriand,
Species described; C. lJaeviy, from Wynard,
Tas, and C, resea, from Pemberton, W.A,
In C. leevis the eggs are pigmented, about
3 mm in diameter, and with discrete capsules-
They are laid in concealed sites on Jand. and
embryonic development is intracapsular. There
are no external gills. After the eges ure flooded
by winter rains the larvae (Fig. 5C) hatch
and undergo aquatic development. The anus
is dextral, The mouth has two upper and three
lower labial tooth rows, and papillae are absent
from the unterior and posterior margins of the
mouth dist (Fig. 6C).
Tn C, rosea the eggs are unpigmented and
have a diameter of 235 mm (Main 1957b),
The entire development takes place on Jand,
and the larva (Fig. 5D) is highly modified,
with no mouth disc (the mouth is a simple
slit), a large yolk sy¢ and an elongate tail, The
anus. is. median,
All members of this species group have one
or other of these modes of development: the
C. Inevis pattern is shared by C. leai and C.
victoriana, and the C. rosea pattern by C’.
lutea (Litlejohn & Martin 1964; Main 1957b,
1963).
CRINIA SIGNIFERA proup
Species examined: C. parinsignifera, C. riparia,
C. signifera, C. sloanei, C.. tasmaniensix,
Species described: C. parinsignifera, from 6
km §.E. of Wandong. Vic.
The eggs are 1.3 mm in diameter, pig-
mented, and with distinct individual capsules.
40 G. F. WATSON and A. A. MARTIN
é 1) Z os
weer
ele 73 2
7s
re pal o
in to
EY Nui ity
“hh v Se Twa a ut ~
wy Sai i
Ag * - # nN aa
h. wun coo tr wl a0 ~
wo ee ec ass
} petra
‘Ry
Mig
oj uy vein in rl fo
ANANTH
Ming
Nu
Won
NAO
Dr.
Aa Ty
: o
Se ange ve)
Ss ne Rowny oN
S Od " eo a)
} A
c
. ~—S
a SS
Pb
‘ m “ai 4 S
ay i me >
yp Ln Se 1 yu yr
Fig. 6. Larval mouth discs of: A, Crinia haswelli; B, Pseudophryne cerroberce: C, Crinia laevis; D.
Crinia parinsignifera; E, Taudactylus diurnus; F, Uperoleia marmorata, In euch case the bar
represents 1 mm,
AUSTRALIAN LEPTODACTYLID FROGS 4i
Ovipasition und developmient are aquatic, Ex-
ternal gills are absent. The Jarva (Fig. 7A) is
unmodified, with a dextral anus. The mouth
has an incomplete papillary border, with both
anierior and posterior gaps, and there are two
upper and three lower tows of labial teeth
(Fig. 6D).
There is little variation in life history in this
group. The same basic pattern is shared by
C. glauerti, C. ingsignifera, C. pseudinsignifera;
C. svbinsignifera and C. tinnula (Main 1957b,
1965; Straughan & Main 1966). In C. fas-
maniensis and C. riparia the eggs are larger
(ovidiameters 1,96 and 2.27 mm _ respec-
tively), and the latter has terrestrial. oviposition
und a lotic type of larva (Martin 1967b; Little-
john & Martin 1965).
Glauertia J overidge
Species described: G. russelli, from Western
Australia.
The cggs have a diameter of about 1.4 mm
atd are laid in water: development is aquatic
(Main 1968). No other details of the life his-
tory have been recorded, and the life history of
G_ miohergt is entirely unknown.
Myobatrachus Schlegel
Species described: M. gouldi, from Western
Australia.
The cggs reach a diameter of 5,5 mm (Wat-
son & Saunders 1959). No other life history
data are on record. but. from the large egg
size and general adult ecology it is very
probable that development takes place on land
(Main 1968).
Myobatrachus is a monotypic genus.
Pseudophryne Fitzinger
Species examined: P. australis, P. bibreni, P.
curiacea, P. correbayee, P. dendyi, P,
remimarmorata.
Species described; P. carreboree, from Mt.
Ginini, A.C.T,
The eggs are pigmented and have a diameter
of about 3 tum. They haye firm, discrete
capsules and are laid on land, in tunnels in
sphagnum. Embryonic development occurs
within the capsule and there are no external
gills. The larvae (Fig. 5B) develop in water.
The anus is dextral and the mouth has two
upper and three lower rows of Jabial teeth.
There are gaps in the papillary border at both
the front and reat margins of the mouth dise
(Fig. 6B).
This pattern of life history—large eggs laid
on land, intracapsular embryonic development,
and aquatic lurval development—is consistent
throughout the genus (Martin 1965, 19674),
with the exception of the Western Australian
P, douglasi, in which oviposition is aquatic
(Main 1964), Blake's (in press) revision of
the myobatrachines includes Meracrinia
nichellsi in Psendophryne, but nothing is
known of its life history (Main 1968),
Taudactylus Straughan & Lee
Species cxamined: 7. diurnus, from ML.
Giotious, Qld.
Ovarian eggs reach 2,2 mm in diameter, but
oviposition and embryonic deyelopment are
not recorded (Straughan & Lee 1966). We
found larvae (Fig. 7B) in a slow-flowing ercek.
The anus is dextral. The mouth structure
(Fig. 6E) is unusual. The jaws are weakly
developed and there are no labial teeth; but
the disc is greatly expanded and umbrells-like,
with a complete papillary border.
The life history of ZT. aeutirostris is not
recorded, and therefore whether or not this
unique Jarval form is typical of the genus is
unknown,
n
wa phdh =) Dagan
meal
Fig, 7. Left lateral view of larvae of: 4. Criniz
parinsignifera; B, Taudactylus diurnus;
C, Uperoleia marmorata, In each case the
bar represents 5 mm,
42 G.
Uperoleia Gray
Species examined: U, marmoratd, U. rugosa.
Species described: U/. marmorata, from 38 km
N. of Bateman’s Bay, N.S.W.
The eggs are pigmented and have discreté
capsules; the ovidiameter is about 1.5 mm.
Development is aquatic, External gills do not
develop. The larva is unspecialized (Fig, 7C);
the anus is dextral. The mouth (Fig. 6F) has
F. WATSON and A, A. MARTIN
two upper and three lower rows of labial teeth,
and gaps in the papillary border at both front
and rear. Moore (1961) incorrectly states that
the papillae extend around the posterior mar-
gin of the mouth disc; and both Moore (1961)
and Lynch (1971) erroneously record that
there is only one upper labial tooth row.
The life history of 0. rugesa is very similar.
Life history characters are summarized in
Table 2.
TABLE 2
Life history characteristics of Australian leptodacrylid genera
Unoper Labia! Gaps in Labial
Egg Mass Development FEatretnal Gills Tooth Rows Papillac
bal
Genus Species 3 Z é = & & 38
Genus Species z= & = 3 Fa E a “4 € =
= a 7 ta w @ = + foe
a = g EN a 5 0 z 2 i=] P< Begs
e zg #4 FF = & > Z << <¢%e
ANELOTUS a ee a=, & ae
LHCHRIODUS + j= -«— oo —-— — + a» JF c=
LIMNODYNASTES ce => r= ~ > —— Le os | — t —
KYARRANLS + = — a ca — + — — + —
PHILORIA ‘ao oe f Eo ie
HELELOPORUS al Ses + = —> ~~ + —F 3 f
MIXOPHYES — + tS fd om @& = 3 «he
NEOBATRACHUS — _ + = = = = = as = =
NOTADEN — & ‘tp 2 ~s =& 4 = ==
CYCLORANA — -b + — 2 — + — —, 7 —
ASSA - ? — y 5 -
CRINIA veoreiana — oo + = 2 = — 77 — le, +:
haswelli — = = = & = 4- = = 4 =_—
faeyis — = t. — — a — + >. — +
itenif = + = + — => _ — No papillae
slenifera sroup — + — — — =
GLAUERTIA a” oe ; = -7 i. = pea
MYOBATRACHUS 2 fen > + >
PSEUDOPHRYNE ~ + 4 a = = = + — — Ea +
TAUDACTYLUS ? oat > nS] = 2 SE
UPEROLEIA >» te br a ae ia | i TS Sion —& 23
Discussion
(i) Status of tHe subfamilies
Life history data support the division of the
Australian leptodactylids into the two sub-
families currently recognized. The Myobat-
rachinae are a close-knit, natural group of
genera sharing several life history features,
These are: eggs with discrete cupsules, ege
masses not foamv; no external gills; « dextral
anus; two upper and three lower rows of labial
ieeth; and anterior and posterior gaps in the
papillary border. The few exceptions are
species of Assa, Crinia and Taudactylus whose
larvae are modified for development in a
parental pouch, or in other specialized niches.
The Cycloraninae are a more heterogeneous
assemblage. Leaving aside C}clorana (which is
discussed. below), there is still a variety of
#Th brood pouch of male
developmental patterns und larval forms in this
group. The frothy egg mass has apparently
evolved at least twice, in view of the occur-
ence of two different methods of foam produc-
tion, In Adelotus, Kyarranus, Lechriodus, Lim-
nodynastes and Philoria the foam is formed
by the female “paddling” with her forelimbs,
which have specialized flanges on one or more
fingers, during amplexus. This paddling causes
a stream of bubbles to pass backward beneath
her body and become entrapped in the mucus
which accompanies extrusion of the eggs (Mar-
tin 1967a). Heleioporuys females Jack these
flanges, and in this genus the foam is. presum-
ably produced by a different (but presently
unknown) method (Martin 1970). Again ex-
cluding Cyclorana, life history features com-
mon to most cycloranines are: eggs with dis-
crete capsules, sametimes in foamy masses!
AUSTRALIAN LEPTODACTYLID FROGS 33
external gills present, 3-6 upper rows of labial
tecth; and no posterior gap in the papillary
border. The anus is usually median, but is
often slighily offset tn Neebdarrachus, and fully
dextral in Adeloms and Mixophyes,
Lynch’s (1971) division of the Cycloran-
inae into tribes is based on breeding biology
and the position of the vomerine teeth. The
Limnodynastini consists of the genera in which
foamy egg masses are produced with the aid
of the Hanged fingers of the female. This group
is of course relatively homogeneous in terms
of life history, since if was partly defined in
this way. The Cycloranini, an the other hand,
exhibit a variety of life history patterns, and
from this point of view do not appear Lo con-
stitute # natural group
(ii) The bearing of life history data on generic
delimitation
The current generic delimitation of Austra-
lian lJeptoductylids is broadly consistent with
what is known of their life histories. In cases
where genera haye very similar life histories,
ea. Neobarrachis and Notaden, Pseudophryne
and the Crinia laevis group, there is sufficient
differentiation in adult morphology and ecolozy
tO warrant generic separation, The retnoval of
Crinia darlingtoni to Assa by Tyler (1972b)
and the subdivision of the remainder of Crinig
by Blake {in press) are supported by life his-
lory evidence. The developmental bialogy of
Assa is unique among Australian leptodacty-
lids, and the Crinia laeviy and C. signifera
groups ure also definable in terms of life his-
tory. The two members of the C. haswellf
group have rather different life history patterns,
but both are distinct from those of the C€-
laevis and ©, signifera groups,
Kyarranus and Philoria are the only genera
whose status seems questionable in the light
of life history data, The similarity between
theny in Most aspects of both adult and Jarval
morphology and ecalogy has already heen
commented on by Littlejohn (19636) and
Braitstrom (1970), and the latter has indi-
cated his intention to synonymise Kyurranus
with Piiloria. Such u change is clearly sup-
ported hy evidence From their life histones,
(ili) The position of Cyclorana
Tyler’s (1972a) contention that Cyclorane
does net cunform with the currently accepted
concept of the Cycloraninas, and has hylid
affinities, is strongly supported by life history
data. Indecd, If regarded solely in lenms of life
history, Cvclorana coincides very clusely with
the pattern typical of Australian bylids ( Martin
& Watson 1971). Characters which it shares
with them, and which are almost unique among
Australian leptodactylids, are the indistinct
egg capsules, the general body form of the
tadpole (particularly the acuminate tajl}, and
the presence of two upper tabial tooth rows
combined with the occurrence of papillae along
the posterior margin of the mouth dise. Data
from other sources, ¢.g, karyotype and mating
cull structure, ure needed hefore a final
decision can be made; but for the present it
shoukl be recognized that the subfamilial dis-
position of Cyclorana and the definition of the
Cycloraninae require revision.
liv) Phylogeny ef the Australian lepioducty-
lids
The phylogenetic relationships of the Aus-
tralian leptodactylids are disputed, Parker
(1940) speculates that the myobatrachines may
have been derived from ihe cycloranines:
whereas Tyler (1972a) regards the myobat-
rachines as the primitive, and the cycloranines
as the denved, group. Lynch (1971) believes
that the two groups are not closely related, aud
that they represent independent descendants
from a primitive leptodactyloid stock,
Our data do not contribute significantly to
resolution of this question. If Cyclorana is left
out of consideration then there are three main
distinguishing features in. the fe histories of
the two subfarnilies. These are (1) the absence
of external gills in the myobatrachines, and
their presence in nearly all cycloranines; (2)
the presence of two upper labial tooth rows in
the myobatrachines, and of three or more in
the cycloranines; and (3) the pap in the lower
labial papillae of the myobatrachines. The
latter two characters stiggest that the
myobatrachines are the more primitive group,
but not necessarily that the cycloranines were
derived From them. The presence of only two
upper labial tooth rows is common in many
families of anurans, ¢,g, most hylids. bufunids
and Neotropical leptodactylids (OQuellman
1970; Martin & Watson 1971; Lynch 1971),
The papillary gap is also a bufonid character-
istic. Thug Lynch's (1971) suggestion that the
myobatrachines may be a relatively unmodi-
fied derivative of the proto-bufonid stock (i.e,
the leptodactyloid group which was ancestral
to the bufonids) seems reasonable, Bufonids
do, however, possess external gills, Life his-
tory data do not assist in the |nterpretation al
cycloranine phylogeny, In terms of life history
characters alone the only conclusions iat can
44 G. F. WATSON and A. A, MARTIN
Key to Genera of Australian Leptodactylid
Laryae
(excluding Glauertia, Myobatrachus)
1. Larvae in brood pouch of adult male -
1. Larvae not im brood pouch of adult Miytniadtardtiaeta:
2, Mouth without labial teeth or papillae... — — — 6. ee
vtssa
.. Crinia laevis group (part)
3
2. Mouth with labial teeth and/or papillae oj... ee ee
Wa aA
. Labial teeth present .
4, Larvae whpiemented; freesuittatoniath i in 2 gtrearits Mt. ‘Glorious # are, + Old.
4. Larvae unpigmented or lightly pigmented
5. Larvae in pools or damp sphagnum,
pherson Range, Qld.
6. Papillac camptetély surrptinding mouth .
6. Papillary border incomplete .... me
7. Papillary border with an anterior gap
7. Papillary border with both anterior and posterior gaps .
8. Mouth with 2/2 labial tooth rows .
8. Mouth with moré than 2/2 labial tooth TOWS ccc ccecee ere cveeene Mh,
9, Mouth with 2/3 labial tooth rows! .....,.
9 Mouth with more than 2/3 labial tooth TOWS ooccce sescceescssuesiusvessesvvecnviveee Oedeomcoeece
10. Mouth with 3/3 labial tooth rows 2.000.006 ee ee ee
10, Mouth with 4-6/3 labial tooth rows _.
. Anus dextral
12, Mouth with 3/3 labial tooth rows —.......-.0...-— ~~
12. Mouth with 2/3 labial tooth rows? —
Labial teeth absent — | occsesisicsrspteerern etree
1. Anus median or neat-median pred aide
4
6
Talay dliurnas
Ris wrist pyTeronegs eel orice tetrocti tute HSE 10000 .. 5
5. Larvae in pools or damp sphagnum; Mt. Baw Baw, Vic, . ~~
or in Hepronipas in earth;
gies Philoria
Point / Lookout, NS.W,; Mac-
; Kyarranus
pes ene Teorhrevrtenetritae! pied ite daeenkirenalaiiattrs Mixophyes
Cyclorana
10
..
~ Holeioporus
Lechriodus
Limnodynastes
Es Adclotus
.. Neobatrachus
Notaden
. Crinia georviana
_. Pseudophryne
Crinia laevis graup (part)
Crinia signifera group
Uperoleia
1 Tadpoles of nearly all Australian Hylidae whose larvae are known also key out in this category.
> Tadpoles of Bufo marinuy also key out in this category.
be drawn are that the cycloramnes afte a more
specialized and less homogeneous group than
the myobatrachines, and do not show any close
affinities with them.
The phylogenetic position of Cyclerana can-
not yet be decided. In terms of life history: it
shows greatest affinity with the hylids, less with
the. myobatrachines, and virtually none with
the cycloranines. It is conceivable that it rep-
resents a relict of a primitive stock which was
ancestral, to both leptodactylids and hylids.
M. J. Tyler (pers. comm,) is currently cogaged
in an snulysis of the affinities of Cyclorana,
and until his work ts completed further specu-
lation is not warranted.
(v) Larval characters as an wid to diagnosis
The larval morphology and biology of most
genera of Australian leptodactylids are
sufficiently distinctive to enuble generic diag-
nosis to be made in terms of these characteris-
tics; they form the basis of the following dicho-
tomous key.
Acknowledgements
Much of the material used in this paper
was collected during field studies supported by
grants from the Nufiicld Foundation and the
Australian Research Grants Committee (Grant
No. 66/16172) to Dr. M.. J, Littlejohn, and by
the University of Melbourne Research Alloca-
tion to the Department of Zoology.
For donating material of species not repre-
sented in our collection we thank Mr. J. M,
de Bavay, Dr. M. J. Littlejohn, Dr. B.S. Low
and Mr. M. J. Tyler.
Some of the illustrations were prepared by
Miss L. M. Howard and Miss M. Leahy,
Dr, M. J. Littlejohn and Mr. M. J, Tyler
read and criticized the manuscript.
AUSTRALIAN LEPTOBACTYLID FROGS 45
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STUDIES ON THE ECOLOGY OF THE AGAMID LIZARD
AMPHIBOLURUS MACULOSUS (MITCHELL)
BY F. J. MITCHELL*
Summary
MITCHELL, F. J., 1973. Studies on the Ecology of the Agamid Lizard Amphibolurus maculosus
(Mitchell). Trans. R. Soc. S. Aust., 97 (1) , 47-76, 28th February, 1973.
Amphibolurus maculosus (Mitchell), the Lake Eyre Dragon, is restricted to arid salinas in the
northern part of South Australia. The lizards live near the margins of the salinas where a suitable
burrow area of fine, wind-blown sand underlies the buckled salt crust. Their essential refuge from
the harsh environment is in the permanently damp sediments, which occur below these dry aeolian
deposits underlying the salt crust.
Preliminary laboratory experiments suggest that this lizard has a high temperature tolerance
(CTM 48.9°C) and low evaporative water losses (1.1 mg/g/hr at 37.5°C)
The harvest ant, Melophorus sp., which occurs in the same habitat, is an important food item for the
lizards, and the elevated rims of the ant nests provide shade and basking sites.
Seasonal emergence, following 3-4 months of hibernation, begins when ground temperature reaches
about 19°C. At this temperature the lizards head-bask, emerging fully from the burrow when body
temperature is raised to about 22°C. Under these conditions the dominant males emerge and
establish territories in mid-August.
Basking postures are adopted to raise body temperature to 37.5°C (eccritic temperature determined
in laboratory tests). In the field, behavioural thermoregulation maintains body temperature at about
39°C during higher air temperatures. If temperature cannot be held down within this range, the
lizard retreats to the humid Jayer below the dry sand in the burrow zone.
Within the population, dominant, sub-dominant and subservient males can be recognised by
behaviour and coloration during the breeding season (September to December). Among the females,
two breeding groups, related to time of hatching, occur. Sperm retention is evident in one of these
groups, in which ovulation occurs about 2 months after fertilization.
Sexual dimorphism in relation to colour pattern is not thought to be the basis of sex recognition
since colour changes occur in both male and female. It seems likely that acute perception of size
difference is the basis of sex recognition.
STUDIES ON THE ECOLOGY OF THE AGAMID LIZARD
AMPHIBOLURUS MACULOSUS (MITCHELL)
by F. J. MircHi.i*
Summary
Mrrcuece, F. J.. 1973. Studies on the Ecology of the Agamia Lizard A.nphiholurus maculosas
(Mitchell). Trans. R. Sov. S$. Aust., 97 (1), 47-76, 28th February, 1973.
Amphibolarits maculasus (Mitchell), the Lake Eyre Dragon, is restricted to arid salinas
in the northern part of South Australia. The lizards live near the margins of the salinas
where a suitable burrow area of fine, wind-blown sand underlies the buckled sali crust.
Their essential refuge from the harsh environment is in the permanently damp sediments
which occur below these dry aeolian deposits underlying the salt crust.
Preliminary laboratory experiments suggest that this lizard has a high temperature
tolerance (CTM 48.9°C) and low evaporative water losses (1.1 mg/g/hr at 37.5"C).
The harvest ant. Melophorus sp., which occurs in the same habitat, is an important food
item for the lizards, and the elevated rims of the ani nests provide shade and basking sites.
Seasonal einergence.follawing 3-4 months of hibernation, begins when ground temperature
reaches about 19°C. At this temperature the lizards head-bask, emerging fully from the
burrow when body temperature is raised to about 22°C. Under these conditions the daminant
males emerge and establish territories in mid-August.
Basking postures are adopted to raise body temperature to 37.5°C Ceccritic temperature
determined in laboratory tests). In the field, behavioural thermoregulation maintains body
temperature at about 39°C during higher air temperalures.. If temperature cannot be held
down within this range, the lizard retreats to the humid layer below the dry sand in the
burrow zone,
Within the population, dominant, sub-dominant and subservient males can he recognised
by behaviour and coloration during the breeding season (September to December). Among
the females, two breeding groups, related to time of hatching, occur. Spetm retention is
evident in one of these groups, in which ovulation occurs about 2 months after fertilization.
Sexual dimorphism in relation to colour pattern is. not thaught to be the basis of sex
recognition since colour changes occur in both male and female. Ut seems likely that acute
perception of size difference is the basis of sex recognition,
CONTENTS
1. Introduction 2.00 2 ee AB (2) Humidity -_ [eee = Cao
4 . 49 (3) Salinity Po «te» no 256
4 Pea ane Metnede ace 7 ag 4. Food and Predation . an L 54
(2) Laboratory data... .. 30 7. Flooding of the Habitat... 58
3. Morphology of Amphibotyrus maculosus 50 8. Activity patterns and Thetmoregutatian 5k
4. Habitat and Distribution a 50 9. Survival in the Preferred Habitat — 4f
The habitat of the Lake Eyre population Tolerance to temperature and water slress 6/1)
a ane peant s 3 10. Establishtnent tnd Defence of Teericory
é burrow zone ..,, Sete ate oth the a Sra be, 62
(3) The “wet” salt zone... .., -. 51 4 A
The Lake Callabonna population... ... $1 11. Reproductive Behaviour ee
The Lake Torrens population - .. 52 12. Sex Recognition ... .., 65
2
Bbyesal panier OF Fie Sasa ie oa 32 13. Growth Rate and Reproductive “Cycle ff
5. Microclimate of the Hubitat ._ 53 !4 Acknowledgements =... 67
(1) Temperature, to oes ae 653) OSL References cor “Erte cto By. See tue “EG
*Late of the South Australian Museum. This paper was compiled from the late F. J. Mitchell's
draft manuscripts and ficld and laboratory notes by Ann Mitchell (Department of Botany, University
of Adelaide. S. Aust. 5001).
4h FE. 3. METCHEL1.
1. Introducton
Araphibolurus maculovas is a highly adapted
lizard, restricted in distribution to the salinas
of the Lake Byre Basin in the north of South
Australia, It was first collected in 1929 by
C. T. Madigan, who lodged with the South
Australian Museum 20 specimens taken at
the southern end of Lake Eyre Nerth in the
area now known as Madigan Gulf. These ear-
less lizards were identified as a variety of
Tympanoeryplis lineata by H. M. Hale, then
Director of the South Australian Museum [in
Madigan 1930). In a revision of Tynpanocryp-
tis, a genus of “earless” dragons, Mitchell
(1948) described the lizard as a new species,
T. maculosa.
Subsequent ostcological studies (Mitchell
1965) showed that the uffinities of this species
lie with the genus Amphibolurus and that it
represents an extreme speciulization within the
venus. The enclosure of the t¥mpanic mem-
brane, which had. led to its assignment to
lympanocrypis, was cansidered likely to be
i secondarily developed Characteristic, This
may be an evolutionary adaptation to burraw-
Ing in dry sand, and may have arisen on more
than one occasion in agamid lizards.
Lake Eyre, occupying an atea of 93500 km,
is the largest of several extensive salinas in the
arid. north of South Australia (Fig. 1), lying
aL the ‘southernmost and lowest end (14 m
below sea level) of the Lake Eyre Basin, This
Basin lies within the 125 mm isohyet in the
driest area of the Australian continent. The
usually dry river systems draining towards
Lake Eyre serve a catchment area of over
1,300,000 kn*. Some of this catchmenr lies
within areas of comparatively heavy und
regular rainfall but only under particular cir-
cumstances does sufficient water reach Lake
Evre to fill the Lake. Following exceptionally
heavy summer rains in western Queensland in
March 1949 and March 1950, Lake Lyre by
September 1950 had filled with water for the
first Lime in living memory. Scientific surveys,
sponsored by the Royal Geographical Society
of Australia (South Australian Branch) and
led by C, W, Bonython, were carried out until
the Lake had again dried out by November
1952 (Bonythan 1955, 1956, L960}.
During this period the lizard was again
observed and collected. [ts ability to spend
long periods foraging on the oper surface of
the salt crust of the Lake, withstanding both
predation und desiccation, aroused oonsiderahle
interest,
In 1964, Donald Campbell successfully
uttempted a land speed record on the surface
of Lake Eyre, In order to gain access to the
solid salt crust, a Causeway was constricted
from the shore-line across the maryinal zone
where the salt crust is thin. The track to the
causeway permits vehicular access to the shore-
line, and the causeway itself, projecting west
from Prescott Point at the tip of Sulphur
Peninsula (Figs. 1, 6), has subsequently
served as a natural “trap” and developed a
population of these hezards suitable for
behavioural studies in their natural environ-
ment. A study area was sett up at the cause-
way, where meteorological observations and
records of hehavioural responses of 2 marked
population of the livardy were made during
visits between February [965 and October
1968.
This. paper iy bused lurgely on field obserya-
tions of the lizard an Like Eyre, together with
briefer studics uf populations on other inland
salitas. These field observations were sup-
potled by vivarium studies at the South Aus-
tralian Muscum, Preliminary experimental
work is reported bnefly.
2. Materials and Methods
(1) Field? Studies
The ecological simplicity of the habitae and
the lack of a predjtor sense itt A. macnlosus
make it an attractive animal for study once
problems of distance, inaccessibility and try-
int werking condilions ate accepted. The
lizards are very alert and hecause of cryptic
coloration almost impossible ta detect except
during their sudden, rapid movements to main-
tain a few metres distance from un approach-
ing observer. During the breeding season,
however, territorial challenges, fighting and
mating continue uninhibited’ by close-range
observation.
Early in the field studies some 10 km of
shoreline were surveyed, but later observations
were resiticled to the causeway population
und all meteorological records reported here
were taken in that area.
In all, L6 trips were made lo the study area
during the period February 1965 to October
1968, Average time spent at the study atea on
these trips Was 7 days. In addition, during
September and October 1967, an extended
survey Was made of the major salinas extend-
ing in an are around the northern end of the
Flinders Ranges. regurded us potentially suit-
able habitats for the Hzards,
ECOLOGY OF AMPHIBOLURUS MACULOSUS 4g
Field data have been derived from several
sources. Continuous temperature records of air
and sand at various depths were obtained using
an EILCO thermistorised twelve-outlet, twin
channel. recording thermometer, while single
readings were taken with a Thermophil elec-
tronic thermometer. Rectal temperatures were
obtained with a Schultheis, quick-reading
mercury thermometer, All instruments were
checked and calibrated in the laboratory before
each trip. Relative humidity was variously
measured with cobalt thiocyanate papers,
Fig.
lithium chloride cell equipment and = whirling
psychrometer.
Behavioural data were obtained from direct
observation and from the marking and re-
Capture of specimens in the causeway popula-
tion, at Prescott Point. Quadrats were marked
out along the edges of the causeway and move-
ments of lizards in relation to these areas were
recorded during the period of study. Lizards
were permanently marked by toe-clipping, and
colour-coded For rapid identification of marked
specimens in the field.
LAKE la
CALLABONNAT=3 *
© MARREE
1. Map showing the major salinas of northern South Australia and the known distribution of
Amphiholurus maculosus. Inset map (upper left) shows portion of Madigan Gulf (the type
logality), and the location of the causeway at Prescott Point which was fhe main study area. A
hlack square indicates areas in which populations’ of A. maculasus have been observed; a white
square indicates arcas where survey of the salina margins revealed no specimens of A_ macn-
losis.
Sh F. J.
(2) Labonuery dare
Experiments on thermal criteria and tem-
perature ltolerunce of A. maculosus were
caried out in an oven preheated to 45°C.
Regular lemperuture increments of 1°C were
made at § minute intervals
Rates of water lass were derived from e¢x-
periments carried out in a thermostatically
controlled, Wauter-jacketed incubator under
temperature conditions controlled to =0,5°C.
‘Yhe test chamber wus also desiccated over
silica gel to ensure a relative humidity of less
than 4% at all temperatures.
Experiments to deterntine water loss through
the integument were undertaken with the vent
of the animal sealed with waterproof adhesive
lupe amd the head inserted through a_ thin
rubber membrane across the mouth of a con-
tainer partly filled wath silica gel to absorh
avy Water lost through the ittegument. The
silica gel was overlain hy u piece of thin card
on which the lizard’s body restect.
The test animals were acclimated for at
least seven days to coriditions involving the
daily allainment of their maximum voluntary
activity temperature (39.8°C), Animals were
tested hetween 1000 and (500 hours during
their period of maximuin activity.
For long-term behavioural studies, a ter-
rarium was set up which successtully simulated
lake shore conditions. A layer of gypseous
clay. maintained in moist condition, was aver-
lain by 15 cm of drv sand from the lake
shore, Over fhe sand, pieces of consolidated
salt crust provided a surface similar to the
burrow zone area.
Both light and heating were provided by
au battery. of high-power incandescent lamps
mounted under an adjustable hood. ‘Time
switches controlled day length, and the posi-
tion of the hood controlled the maximum
temperature, Strong correlation between field
observations and laboratory tecords was Found
for daily and seasonal activity cycles and the
onser of reproductive activities.
3, Morphology of Amphibolurus macplosus
Amphibolurus maculesus shows several
features OF morphology, physiology and be-
haviour Which reftect its adaptation to arid
silinas, Its dorsal surface is white to very
pale grey with u row of very dense black
blotches on either side of the vertehral line,
Dorsally and laterally there are smaller black
and rusty-hrown pigmented areas, This colour
MITCHELL
pattern gives excellent cryptic coluyralion on
the salt surface, the dark areas blending with
the shadows of small holes and pinnacks on
the rough salt (Figs. 9, 10, 15). The ventral
wurfuce is white with a dark streak cxtending
longitudinally along the centre of the throat
to the gular fold, Colour variations related to
the environment, ani the striking colour pat-
terns developed during the breeding season
will be discussed later in this paper.
The adult male attains a total length of
about L1.5 cm (snout-vent length 7 cm}, the
adult female being smaller ut about 10.0 cm
(snout-vent length 6 cm).
The eye ts small and deeply sunken and is
protected from salt glare by promiment ser-
rated eyelids with dark pigmented Jinings
(Fig. 4), The nostrils, while showing a circular
external opening, open inte the nasal cavity by
only a narrow slit, directed forward and down-
ward, This structure prevents the nasal pas-
sages from becoming blocked when the lizard
burruws through joose sand. Speci! nasal
stwucture has been described by Stebbins
(1943) in the Americun venus U/ma, which is
a sand-burrowing form. He has shown (Steb-
bins 1948) that other American iguanil
genera including Uru, Helbroakia and
Phrynesonea, all of which habitually burrow
in loose sand. have similarly adapted nasal
structure.
4, Habitat and Distribution
The type locality of 4. meaculasies 1s Madi-
gan Gulf, Lake Eyre North, out from the
mouth of the Frome River. The species is
widely distributed around the southern shore-
line of Lake Eyre North and around the
nerthern ind south-evstern shorelines af Lake
Eyre South. Throughout the study every oppor-
tunity was taken ta search other lakes for
additional populations and practically all of
the major salinasy in che State were at Jeast
cursorily cxamined for signs of habitation by
these lizards. Two additional populations
were found: uw population in the saline areas
at the southern end of Lake Callabonna and
extending soulhwards through the salt chan-
nels inte the northern tip of Lake Frome, and
nother population along the north-western
shoreline of Lake Torrens, Fig. 1 shows the
area searched, the localities from which speci-
mens have been taken, and the range over
which this species is known to occur.
The geomorphology and the history of land-
forms In the Lake Eyre Busin have been dis-
cussed by Wopfner & Twidale (1967). Evi-
ECOLOGY OF AMPINBOLERUS MACULOSUS 3|
dence from fossil deposits suggest that exten-
sive areas Of brackish water or saline swamps
existed Within the basin from mid-Tertiary to
the end of the Pleistocene. The ongin of the
salt in Lake Eyre has. been subject to debate
but i stems likely that both cyclic sult and
connate salt, accumulated from waters brought
inte the basin from the extensive drainage
system, have contributed (Bonython 19546;
Wopiner & Twidale 1967; Twidule 1968),
The present isolated populations of A, mracy-
fases, which although spatially widely sepa-
rated ure virtually morphologicylly — indis-
tinguishable from ane another, may be relict
from uw more widespread species evolved in
association with the shoredines of the Sarge
brackish lnkes and swamps that were a feature
of the Lake Eyre Basin during Pleistocene and
early Recent periods.
THE HABITAT OF THE LAKE EYRE
POPULATION
The habitat within the study area can be
divided into three distinct zones, the “beach”.
the “burrow zone” and the “wet salt zone”
(Fig. 5). This zonation can be readily identi-
fred throughout the distribution of the Lake
Eyre population examined, bit is much less
distinct in the area of ibe other two poepttly-
tions.
1. The beach; tn tie area along the southern
shoreline of Lake Eyre North which was
most intensively studied, the beuch is backed
by near-white sandhills varying from low
consolidated hummocks stabilised by low
shrobby vegetution { Nitraria, Seaevola) to high
drifting dunes encrouching upon the lake. In
Many arcas low cliffs varying in height from
about a metre to upproaimately 12 m above
the lake hed expose Recent to Cretaccous sedi-
ments to face the lake Continuous wind
erosion transports large quantities of gypsi-
ferous clay and grit and other aeolian fines
out over the beach and on to the surface af
the lake, The beach is usually a narrow strip
of consolidated sand Frequently intermixed
with wreas of very coarse sand and pebbles.
In areas where it is not overlain by recent
drift it remains damp during the cooler months.
Typical pebble beaches occur in some areas
and the height and form of some of these
beaches suggest sustained periods of strong
wave action in the very recent past,
2, The burrow zone. This term has been
adopted to cover the zone tn which most of
the activity of the lizards takes place. It con-
sists Of un trea of distomed, dry, crusty sur-
face, varying From a few metres wide (im areas
where flooding upproaches close inshore or
where little pypsiferous drift is brought out by
the prevailing wind) to 400-500 m wide in
stidom-flooded embayments where wind-
drifled deposits have accumulated on the sur-
face (Fig. 5), This 5-15 em thick layer of
fine gypsiferous clay and sand rapidly dries
out and in doing so cxpands and distorts the
Salty crust which Sorms over the top of i.
In section, this provides a burrowing substrate
consisting of an irregular, salt-impregnated.
crust up to 2 em thick, lying aboye approxi-
mately 10 em of fine, dry, windblown sedi
ments underlain by sand which is kept con-
tinvally dump from the water-table 40-70 em
below,
The lizards break through the salty crust at
a weak point and then literally “swim” through
the fine mobile scdiments below until they
encounter the high humidity associated with
the damp consolidated sand at lower levels.
They remain there until their re-emergence js
triggered hy rising temperatures or some
internal stimulus.. They follow the thermal
gradient hack to the surface, frequently
emerging at a different point to that of entry,
3. The “wet" salt zone: The solid salt crust of
the lake varies from about 2 cm in thickness
in inshore areas to about 40 cm in thickiess
fowards the centre of Madigun Gulf. This
crust, overlying wet gypseous clay. becomes
distorted and buckled as the salt recrystallises
on drying out (Figs, 7, $). Although most of
the activity of this lizard takes place within
250 in of the shore, both the lizards and the
nesta of the ant Melopkorus sp. (their princi-
pal item of food) have been observed more
than 1400 m from the shore linc, During the
territorial season, many young subordinate
mules are forced to live outside the burrow
sone and are frequently found well offshore.
While liltle weight can be placed on the salt
surface without bringing water to the surface,
the actual surface “skin” js dry most of the
lime except in cerlain effiorescent areas close
to shore which are continually damp. Relative
humidity readings using cobalt thiocyanate
papers set | cm off the salt under a shield.
gave readings as high as 55% over these
efflorescent areas, bul in other areas the read-
ing was below 15% and equalled the reading
at 100 cm above the surface.
THE LAKE CALLABONNA POPULATION
Essentially the same conditions prevail as at
Lake Eyre. with the lizards living around the
52 Fr. J. MITCHELL.
southern margins of the lake on isolated
patches of aeolian drift in association with dry
saline crust, A suitable habitat is not present
in northern parts of the lake, where fresh
water from the Strzelecki Creek has leached
the bulk of the soditim ancl catefum salts from
the surface deposits and the water table is
deeper hecause of extensive Recent lacustrine
deposits on the surfuce.
THE LAKE TORRENS POPULATION
The surface of Lake Torrens is wet and
muddy und the take frequently contains water.
As a consequence the population has adjusted
its behaviout to living amung low vegetation
along the immediate shoreline and upon jow-
relieY islands covered with chenupodiaceous
shiulbs.
PHYSICAL FEATURES OF THE BURROW
ZONE
Field and laboratory observations suggest
that there are three essential habitat charac-
teristics Which influence the cistribution of
this lizard,
|, A surface crust: Specimens placed in a
laboratory cage containing either loose sand ur
consolidated sand lacking a surface crust were
upable ta burrow, ‘They were only able to
scoop oul a shallow depression, Several ficld
observations related to this were made during
the 1952 flooding of the lake when most of
the lizards were forced onto the damp beach.
Some did succeed in finding crusted areas back
among the veyetation but several lizards were
found im shallow depressions against drift-
wood wong the beach. Presumably the sur-
fave crust is casential in providing initial pur-
chase against which these lizards start their
“swimming” procedure in burrowing through
the sand, Vhe consolidated surface crust need
no. Hecessarily be saline and in fuct, gypsiferous
sediments hus Logether by filamentous blue-
green algae may well have provided a suitable
hafritat surface over wide areas in the past
and may prove io be aw contributitig factor at
present.
2, A layer of fine dry sand (and/or clay) about
10 em thlek under the erust: In order to pro-
vide adequate insulation from the extreme
heat of the summer, the fine sand or sand«
yypsiferous clay mixture must be dry and
about 10. em thick. It is probably important
that the sediments contain pypsum to promote
expansion and rapid drying after wetting.
3. A constant source of humidity Jor Me re-
treat: Because of the water balance problems
inherent in living upon a salt surface sub-
ject to exlrerne lemperature, mw is essential
that the environment provide a refuge in which
both evaporalive und respiratory water Josses
be minimised during the periods of retirement
underground. The sediments al about I) cm
helow the surface crust must be permanently
damp,
PRESENT KNOWN DISTRIBUTION
At most of the localities examined along
the shore of Lake Eyre North and Lake Fyre
South, lizards for “signs of their presence)
were observed whenever a combingtion of the
three factors described above were found. This
applied equally well to Lake Callabonna ex-
cept that suitable habitat there is much more
limited. Most of the northern part of Lake
Callabonna and Lakes Blanche and Gregory
have uo very friable surface deposit of fine
gvpsiferous clay which is not bound hy a
surface crust. This is not a suitable habitat.
Lake Torrens lies on the “weather” sie of
the Flinders Ranges and is therefore subject to
more frequent flooding than the other lakes
around the northern end of these ranges. The
lake surface is therefore permanently wet and
does not provide a suitable habitat, but at the
northern end and particularly along the western
side, suitable conditions occur along the actual
shoreline and upon Jow insular greas where
the pale gypsiferous clay supports sparse
halophytic vegetation, Many of these arens dry
out during the warmer months, and form a
suitable habitat with a distinct surface crust
underlain by varying depths of dry pale yellow
to red gypsiferous clayey silt, This habitat i
exploited by the Lake Torrens population,
Specimens have been collected up to 275
metres back into the vegetated zone away
from the margin of the lake,
Chher Jakes examined were Lake Harris,
Lake Gairdner, Lake Everard jind Lake Hart
(Fig. 1). Although knowledge of the shore-
line conditions gained during this survey was
very fragmentary, the yeneral information
gathered is considered adequate lo suggest
that if any additional populations are found
on any of these lakes, they will not be found
living upon the surface of the lakes as at Lakes
Eyre and Callabonns, but along the margins
as at Lake Torrens. The sufface conditions
of these Lakes ure similar to Lake Torrens,
the surface being too wet le support the ants
ECOLOGY OF AMPHIBOLURUS MACULOSUS 52
(Melophorus sp.) which provide the principal
item of diet for these lizards; these ants, or
other species of similar burrowing habit, are
restricted (a the shore-line areas. In many of
the Jakes, the ecological situation occupied by
the unis ut Lake Eyre is taken over by 4
species of Fighting Spider (Geolycosa sp.).
Cicindeline beetles are also frequent and active
predators over these damp lakes and in part
Teplace the ams as surface scavengers, They
dig vertical burrews into the damp muddy
surface an@ huve only been seen ut the Lake
Eyre stiidy area when layers of fresh silt from
fioodjags were overlying the salt:
5. Microclimate of the Habitat
A. macilosus has a strict preference for the
Taargins of the salinas and does not invade
the white sandhills despite ts competitive
daminunce., in cage experiments, over the
only agamit species which occurs there
(Amphibolurus pictus Pevers), It was noted
also that 4. pictus makes burrows over a
metre Jong and up to 20 cm deep in mid-
summer, despite the fact that its thermal
tolerances are only slightly inferior ta those of
A, maculosus (see Table 2), This suggested
that the lake surface environment may have
some thermal advantage over the adjacent
sandhills, Climatic conditions prevailing within
the habitat were studied in an effort to deter-
mine the reason for the strict habitak prefer-
ence and also to provide a foundation for an
analysis of the activity and behavioural pat-
terms observed,
(1) Temperature
Using the thermistorised recording ther-
mometer, aa allempt was made to compare
the temperature profiles of the sections. of
the burrow zone preferred as a refuge by
the lizards with parallel situations in the
adjacent sandhills. Comparison between sum-
mer (February) and winter (July) temper-
alure patterns for both the burrow zone area
and the nearby white sand-dune area js given
in Fig. 2. Summer soil temperature profiles
for burrow zone and sand-dunes aré shown
in Fig. 3, The data obtained proved difficult
to evaluate, probably because of the difficulty
of obtaining strictly comparable test. sites, The
ground humidity varies from place to place
with the local soil structure conditions and
the thickness of the surface drift, and the
extent to which the ground water has pene-
trated towards the stirface also varies. It is
nut possible to assess these fuctors without
breaking the surface crust and thereby dis-
turbing the stratification of the protective
layers winder which the lizards normally rest.
Therefore, while the lizards were found con-
aistently to prefer to rest on or just in the
damp sand underlying the mobile dry sand,
it Was seldom possible to determine wher
the tip of the thermisior probe was lying in
this same position. Aljso the protective value
of the surface soil was greally reduced by
local #ain which increased its conductivity, All
of these factors reduce the comparative value
of the data.
"The data, however, do allow iwo conclusions
to be drawn with reasonuble confidence.
Firstly, during the cooler weather when both
the burraw zone deposits and the adjacent
sandhills are damp to the surface, the thermal
characteristics are essentially the same. Second.
ly, in midsummer the temperature at average
refige depth (10 cm) in the burow zone,
probably due to the thermal capacity of
the water table 40-70 em below, is lower
und subject to less fluctatlon (30=1,5"C)
than the temperature in an equivalent situation
in the adjacent sandhills (34%4°C) (Figs.
2, 3).
{2) Humidity
Surface temperatures were recorded using
a contact thermistor and a Thermephil elec-
tronic thermometer. Because of the salurution
of the lake floor right to the surface and the
periodic appearance of free water on the sur-
face of the more thickly salt encrusted area,
it was anticipated that evaporative cooling
would contribute to lowering sutface temper-
ature of the Jake and raising humidity, particu-
larly at the “living level" of the lizards (1 cm),
both being features which would improve its
suitability, as a habitat during the warmer
months. However, careful testing with the
thermistor probe revealed the acttial surface
of the “wet” salt to be dry and at the same
temperature as the crust in the burrow zone
and ue sand in the sandhills. Similarly, tests
with cobalt thiocyanate humidity papers at
1 cm and 100 em above the lake surface
and 100 em above the adjacent sandhill sur-
face usually gave the same treading, Over the
temperature range 30 to 40°C, during which
lizards coukl he expected to be active over the
“wet” salt surface, the readings were usually
below the minimum sensitivity of the paper,
mdicating relative humidities below 15%.
a4 F. J. METCHELL
Humidity teadings, derived from each af
the sources described ubove, varied from Jess
than 5% to 55% refative humidity. ‘The
higher readings invariably followed light rain
or thundery conditions and were the same
for sandhills and lake surface sites, As air
lemperuture increases each day, over the solid
salt crust, the “wet” salt begins to “sweat”
und brine appears at the tips of small, self-
sealing, salt pinnacles. Due to the high tem-
peratures and high evapurative rate it ts likely
that. any humidifying effect of this brine is
restricted to a microtayer, of perhaps only
a few mm, just above the evaporative sur-
face, However, tf was not possible to detect
higher humidity over the “wet” salt surface
and this is possibly due to inadequate instru-
mentation,
13) Saléstey
The salinity: of the wind-blown silts deposi-
ted in the burrow zone varies with depth.
Salts dissulved out from the superficial layers
were 7-11% of the silt; 3-5 em down, 4-8%
and 5-10 em down, 3-4%. Below this level
the salinity steadily increases again up lo
saturation at the water table 40-70 cm below
the surfate, The decrease in salinity nearer
the suftface is probably due to the recent
ofigity of the surface deposit and to condensa-
tion dlong the line of demarcation between
the permanently dampened sands and the
overlying dry sediments leaching the salt out
over a period of time. The lizard} normally
rest in this zone of lower salinity but whether
this is of any advantage is: unknown. The
sand along the foreshore also contdins 2-4%
salt and the observed salinity stratification may
he due to an increuse in surface salinity hy
wind drift from the sandhills rather than to
desalination of the intermedinte luyers.
Several small areas of white efflorescent
salts were found just outside the burraw zone
and these areas are continually Josing water
la the atmosphere, Although no. hzards were
ever ubserved using one of these areas on a
hot day. the large number of fecal pellets
Fig.
present on and around them suggest that their
thermal sdvantages may be ulilised by the
lizards. The surface temperature of such an
area of about 3 m* near the main study site
was compared with that of the surrounding
salt and found to be up to 9.5°C lower
It appeurs that the Jake surface confers no
advantages upon an inhabitent, either in terms
of surface temperature or humidity, over the
condition prevailing in the surrounding sand-
hills. Despite the abundance of free saline.
water throughout the habitat it may still be
inadequate to prevent the high evuporulive
rate [about 220 cm per annum (Bonythen
1955)] from creating sufficient dry hygroscopic
salt throughout the habitat to place greater
strain on the water balance of this species
than other desert-adapted lizards.
The permanently damp layer under the pro-
teelive crust in the burrow zone provides. a
high humidity retreat into which the hzards
ean Tetire overnight or, if under social or
environmental stress, for much longer periods
without endangering thew water balance. This
is regarded as the key factor an restricting
the species Wy the marginal urcas of the
salinas.
6. Food and Predation
The main food source for A, macilosus is
the harvest-ant, Melophorus sp. In addition
to providing food, the nest-mounds of these
wots are important “features” in the generally
featureless habitat, providing lookout points,
basking sites and the only source of shacle.
Colonies of Melophorus generally occur on
the “wet-salt” surface usually within 750 mm
from the beach zone; preference is shown for
areas where the salt-crust is no more than
2 em thick and underlain by telatively clean
sund down Lo the water-table 40-70 em below.
The colonies are regularly spaced over the
tuke surface, each being about 10 m equi-
distant from the next (Mig. 8}, Investigation
of a nest revealed a setics of upper gsuallerics
just beneath the sult crust and i single verticy!
hole z0ing down about 40 ein to a second set
2, Comparison between summer and winter thermal characteristics of the borrow zone and adja-
cent While sand dunes, Figures from continuous recorder traces of temperature were platted ut
40 minute intervals.
~ temperature at -10 em
-——--—- lomperuture at -l cm
-- +). oir temperature at 40 em above ground
4A) Burrow zone temperatures over 34 hour periods in February, 1965 and July, 1966, Tem-
Peroture range at--10 cm for the February period was 3.5°C, about a nivan of 33.6°C.
(B) Sand dune temperatures. over 34 howe perinds in March, 1965 and Tuly, 1966. Temperao-
thre range al -10 em for the March period was 1 1L.5°C. about 4 mean of 34.6°C.
ECOLOGY OF AMPHIBOLURUS MACULOSUS 55
A pe Pky
7 ead! xt teal
é ~ o
c N ¢
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| ¢ \ 7
4 LN eo
-40 r “4s Loe
u fy ut Re fat
é a tan
L < Le
/ —T te ee : f
¢ a yo vt
ay as mee ee ee
! re fy lark 2
E ay 22% FEBRUARY 1965 Ore cee ae an aad
M~30 oF he Pee . :
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E-s Ls s woot ‘
. a Rare
t
U;
R Fete
Ee - a.
oAP20 er ate f Ls
é i ra ON
/
{ sf
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¢
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ee
19 JULY 1964 har a 20"
é 8 10 12 14 16 18 20 22 24 2 4 6 8 10 12 14
ee ee Se ee a ee! ee ee oe es en re” jee i L =| = ! | fe D Re | homeo, |
TIME - HOURS
| 40
2°49 MARCH 1965
T
E
ME
P
E
R
A
T “7S
U e ie
R
PY te
°C
23" JULY 1966 Tee ee sry Pull 2ath
HHO ys Tee
6 8 10 12 14 16 18 20 22 24 2 4 é 8 10 12 14
be L i i He ees 1 L ni Ly gt AP estoy ke L be Ves L 4
TIME - HOURS
FIG. 2
SH FJ. MITCHELL
of galleries at or slong a consolidated layer
of clay just above the water-table, The Gon-
suruction of these galleries produces, on the
surfuce, a distinctive crateriform mound up
to 2) cm ja height. The sand below the salt
crast is usually covered by a fine layer of clay
Which probably accounts for the reddish-
brown debris, brought from below, being in
marked contrast to the while salt crust,
At any time, only 9 small proportion cf
the total wumber of nests contain active
colonies, The ants show linited activity dur-
ing the day, appearing at the surface only to
deposi sand grains from the workings below
round,
In yicw of the limited aboye-surface activity
of the wnts, it has been suggested (Madigan
1930) that Melophorus baryests micro-
organisms either in the salt crust (blue-green
algal cells) or in the damp sand above the
water table (Dunalielle spp.). The nests usually,
however. contain insect remains. and the ants
have been observed gathering bodies of insects
trapped on the sult. Probably most foraging
uclivily by the ants oecurs at night.
A. niculosus, also, is an Opportunistic
feeder and when other insect life is available
on the lake surface, the lizards feed on a wide
ranve of different insect species. On most
nights numbers of insects are stranded on the
salt. surface. Their presence may be explained
in several ways, With jin, offshore wind many
insects may be blown out over the lake;
others may be attracted by the rise in humidity
over the lake surface af night; aquatic insects
(particularly on moonlit nights) may mis-
lake the white lake surtuce for walter. Overall
the Jake surface produces a vast “white sheet”
effest — un cimmmonty used form of insect
trap—and many insects are Ifapped on the
hygroscopic salt surface and killed when
temperature increases next morning, During
ihe spring months, September—November.
vast quantities af insect Jile may occasionally
be stranded on the lake surface. The bodies
of insects Which have been neither captured
by 4... mueu/osny nor later scavenged by Melo-
hig,
phorus become incorporated within the sur-
face salt layers.
it ls Of Intevest thal the characteristic insect
fauna developed in the vegetation of the shore-
line and sand-Aills is rarcly stranded on the
salt surfuce of the like, while the majority
of species stranded on the salt are seldom
seen about the shore.
The period of greatest feeding activity of
A, maculasus (February to April) frequently
coincides with drought conditlons in the sur-
rounding country, and consequent limited
supply of windborne insect life.
Tn view of the limited number of ants active
on the surface, the lizards nist lorage out
over the lake surface for long periods at high
temperatures. The lizards dig into the sides
of the ant nests to retrieve ants from inside,
and the more frequently visite ant-nests
closer tnshore become very battered in appear-
ance, Shallow depressions scraped out at the
base of the nests provide small areas of shale
intu which the lizards may retreat.
The Lake Dyre population ol A. maciulosus
displays no response to overhead predators,
relying lor safety on their cryptic coloration,
The only possible protective cover on the lake
surfuce ts provided by buckles and cracks in
the surface of the salt crust (Figs, 7, 13).
Although these would seem to provide ideal
reweuts they arc tarcly used by the lizards.
The crystalline swit in ihe erust provides a
“glasshouse” eflect and the temperature below
the ¢mist may be several degrees higher than
the surface of the salt. When pursued aver
the surtace until exhausted, the lizards are
more likely to come to rest in the shade of
like pursuer than to retreat helow the salt crust.
There is Jittle evidence of other potentiil
Predulors moving ot across the salt surface,
Hawks. constitute one Of the main predators
of A. piceus from the sandhills adjoining the
Jake. On several ovcusions dead specimens of
A. picts have been observed well off-shore
on the fluke surface. Presumubly these have
been dropped by hawks. The fact that these
3, Yautechrones (soil-air temperature profiles at. different times during the day) recorded on 6th
March, 1965 for (A) burrow zone area and (B) sand dune urea. Differing thetmal characterts-
tics of the two localitics result in the more rapid and greater heating in the sand dune area,
At—I0 cm (averave burrow depth of A. muculases) in the burrow zone there is a temperature
range from 30,9 to 35.6°C over the timc period recorded. At the same depth in the sand dune
area there is a temperatitre ratige from 31.4 to 4L,.4°C over a similar time Interval, In (B), due
to the limiled range of the thermometer used (calibrated up ia (20 F (49°C)), no figures are
avallable for (he 2 em and 4 em depths at ihe 1120 and 1515 readings. Field notes recorded
that, at these times and depths, the temperature Was “12Q°F-+> nd rising rapidly”.
ECOLOGY OF AMPHIBOLURUS MACULOSUS
4 se } ¥4
57
SURFACE
~1
2
DL
E |}
E 7 0630 1100 1600
H
25 30 35 40 45 50
— 1 1 aoa 2 —- lhe sie, ® re 1 Sy
1“. 1
TEMPERATURE ‘°C
i> Z \ \
SURFACE + - = a aa si
|} I as 5
0600 Q?30
TIaAymg
—1
N
3
po
pe)
10 ? 1120 1815
25 30 35 40 45 50
t . i P af t
TEMPERATURE ‘°C
FIG. 3
38 BF. J. MITCHELL
captures have not been retrieved would sug-
west that the glate From the Jake surface may
confuse werial predators.
7.. Flooding of the Habitat
The periodic flooding, either of the whole
ot Luke Eyre or, following more localised
ruins, of restricted areas, must affect both
Melophorus and A. maculasus:
Melophorus colonies withstand flooding for
consideruble periods by becoming sealed off
below the water. As water disappears from
the lake surface, the colonies soon “break
out” wand tework the galleries and nest. The
shape of the above-sutface mound would give
protection to the opening of the nest during
light flooding.
When the nurmal burrow zone is Mooded,
A. maculosus retreats to the shore line and
lives, precariously, in the sand there. The
lizards are frustrated in atlempls to retire
underground since there is no consolidated
crust. to provide Jevernge for burrowing, and
they have no refuge from predators such as
the Ground Goanna {Mararnus gouldii Gray)
ugainst which they have adequate protection
in their narmal habitat, Among the lizards
forced ashore by flooding, colour changes
occur in which the backgtound colour of the
lizard becormes a deeper grey and the areas of
black and rusty-brown pigmentition become
more numicrous and intense, giving 2
speckled appearance more similar to the pebbly
heach sands, At times of flooding, large nuim-
bers of Silver Gulls (Larus novae-hoflandine
Stephens) ure attracted to the arca and these
probably constitute the greatest predation
hazand tor A- maculovay.
An interesting response 10 flonded conditions
has been observed in A. maculosns, During
winter at fow ground temperatures (down to
16°C) the lizard is in torpor. In vivarium
experiments at low ground temperatures, (14-
16°C}, flooding of the substrate aroused lizards
which had been belew the surface in torpor
for 3-4 weeks. Emergence to the surface
involved a greater degree of muscular ¢o-
ordination than is usually shown at these
temperatures nnd may be triggered by lack of
oxygen. Inflation of the body when it emerges
from underground ensures that the lizard floats
on the surface of the water. It would appear
possible that such a reaction would assist
survival of the animal in face of encroaching
floadwatets.
Localised areas of water in the lake show
marked response to changes of direction and
velocity of wind, A flooded inshure area may,
within a few hours, appear us an exposed
sull surface anil vice versa, This dust impose
uidditional problems in the re-establishment of
both Afelapherus and A. maevloases in their
preferred habitat,
8, Activity patterns and Thermoregulation
Like other ectothermic animals, the activity
af Amplibolnvus macelesus depends on
temperature. The lizards show definite be-
havioural fesponses al cerlain temperatures
and these responses serve to keep the body
temperature within a preferred activity range.
Throughout the season of lizard activity. varta-
tions occur in the means and ranges of these
temperatures, These variations, imposed by the
uge and sex of the particular lizard and its
position in the social structure of the popula-
tion, are considered later, The basic range of
temperatures associated with various wetovities.
summatised in Table 1, fs dictissed here
During the winter months (May to Augusr)-
A. maculosasy remains in torpor helaw ground
Emergence threshold air temperature is 19°C,
at which temperature there is early season
emergence (August-September). under stress
of hunger and drive for the establishment of
terrttory, in males. Ac this temperature only
the head of the lizard is thrust through the
salt crust. It is darkly pigmented in the pineal
tegion and the eyelids are extended, indicat-
ing accumulation of blood in the cephalic
sinuses, The morphology of venous shunts in
the cephalic region, and their role in head-
hody temperature regulation, have been
described for Pkrvnetame cornutum by Heath
(1964, 1966), Although the anatomy of the
head of A. mtack/osus has not been examined,
there is similarity in head-basking hehaviour
hetween this lizard and PArynesoma, Which
is Known to use this aid to rapid stimulation
of the central nervous system and elevation
of body temperature. It seems likely that A
maculosus usts a similar mechanism. The
head-busking position is only maintained until
the body temperature is raised to 22°C, at
which temperature the body is Fully exposed
at the mouth of the burrow, Tt is at this tem-
perature that most lizards first emerge for
the season in September—October. During
early busking stages the body of the hzard
is orientated with the back cxposed to the
sun (or source of heat in the vivarium), and
the body ts inflated laterally and closely
ECOLOGY OF AMWPAIBOLURUS MACULOSUS ay
pressed to the substrate, with the hind legs
spread out flat behind the body in a “swim-
ming” position,
Under laboratory conditions, with air tem-
perature at 23°C, the adult lizards emerged,
with eyes still closed, as soon as the lights
came on. Within 10 minutes all were fully
emerged and awake in the flattened basking
position. Within 30 minutes rectal tempera-
tures had risen to 30°C at an air temperature
of 23,9°C,
During summer, mean burrow temperature
(al about 10 em depth) is 32°C. Under these
conditions daily emergence occurs with first
light, and temperature is no longer the cons
trolling factor.
The basking position is maintained until
body temperature reaches 26-27°C, when the
hzards first start to move away from the
burrow and ate sufficiently co-ordinated to
feed.
At a body temperature of 32°C, the lizards
ure capable of Cull range of movement and
activity and, in hot weather, they move out
onto the salt crust at this temperature. The
eceritic or preferred body temperature, was
determined in the laboratory by use of a saw-
tooth thermocline. Under laboratory condi-
TABLE I
dency of A, maculosus in relation te body
femtperature
j
| |
|
ee |
49,5 °C tothal temperawre uncontratud
=48.9°C—CTM f hentine
|
aso!
|
2-43" C omantenee
aA = | temorceutatory
40°C - tem neruture
—39C—oominiled Lemineraure (lod) | enntan|
AIS ervenitictonierature Cab,)
55° |
| 26¢>—nn Activity
eae
27-289 C—maye frm burriy Huskie range
ais Oe
12° C—ful emergande
19% —rarly emergence; head basking
tions the top temperature of the basking range
is 37.5°C. At this temperature there is re-
orientation of the lizard from the positive to
negative with respect to heat source,
In the Vivarium, the lizards can retreat
under-grournd at any time, whereas in the lake
habitat, they must return from the wet sali
surface ta the burrow zene to retire under-
ground, In the vivarium they retreat under-
ground at a body temperature of 37-37,5"C,
While in the field they remain active on the
salt surface away from the burrow zone with
body temperature held at about 39°C.
Except under very hot conditions, the lizards
have a normal activity cycle of about 8 hours
per day. This long period of scavenging is
correlated with food supply and is necessary
lo ensure that they obtain sufficient food. In
summer, during, much of this time air and
surface temperatures would be over 45°C. At
these temperatures the body of the lizard is
held high off the salt surface, the only points
of contact with the bot surface being the
feet and end of the tail. If stationary for any
length of time, the toes of the hind feet are
Taised from the surface so that there is con-
tact only with the limited area of the heel.
During thermoregulatory maintenance of
hody temperature at ahout 39°C, use ts made
of the diversity of substrate conditions pro-
vided by the buckled salt crust, of the small
ateas Of shade cast by the ant nests and of the
few small pieces of driftwood embedded in
the salt surface which ate used as elevated
perches,
Since, during the hotter summer months.
there is an established behavioural pattern
of daily movement out from the burrow zone
onto the “wet” salt crust as the temperature
tises, it seems likely that the lizards achieve
same thetmoregulatory advantage in this
ured.
While measurements revealed no consistent
decrease of temperature or humidity over the
“wet” salt, this may simply reflect the techni-
cal difficulty of detecting small changes on the
actual surface of the crust.
Slight pressure on the “wet” salt crust
releases moisture to the surface, accompanied
by 4 rapid, transitory fall in temperature due
to evaporative cooling (particularly when a
wind is blowing). The lizards, by scratching
at the ‘surface of the crust, may achieve this
same cooling effect for parts of the body in
contact with the surface, It is noticeable that
AD TP. J. MITORree.
during the hot months, which coincide with
the defence of territury by the dominant
mules, the young males forced to spend con-
siderable time out over the salt surface devel-
op uccumulations of salt on the digits of all
limbs (Vig. 11). There may also be salt
encrustation along the ventral surfaces of
these lizards. This sale accumulation may be
caused by the lizards seratching at the sur-
face of the salt crust in allempts to bencfic
from any slighl evaporative cooling elects.
When no longer able to control body tem-
perature by adjusting position and stance, the
lizard Must retreat fo the burrow zone and
retire underground, In order ta reach the
cooler, wet sand Jayer near the water table.
the lizard must first penetrate the Joose sand
below the salt crust, The upper layer of this
loose sond muy reach w temperature of 60°C.
A, maeulosus must have a short-term toleranee
to this extreme temperature in order to benefit
from the hwnid layer below,
Wf Foreed to remain active at higher temper-
atures, A, wraculosus begins to “pont” at a
hody temperature of 42-42.5°C. In this action
the mouth is opened wide (Fig. 10), and
there is u steady rate of deep breathing at 60
breaths/ minute. Unlike some other lizards
(ew, Varanes spp. A, pietus, A. tnermts)
there arc no associated movements of the
gular pouch during panting in A. muculosts,
Tolerance of temperatures. above 42°C, at
which thermoregulation begins to break down,
are discussed below.
Preferred body temperatures and thermo-
regulatory behaviour of several other species
of Amphiholuruy Wave been described by
Bartholomew & Tucker (1963). Bradshaw &
Main (1968), Brattstrom (1971), Heatawole
(1970), Lee & Rarham (1963). Licht, Daw-
son, Shoenwker & Main (L960), and Pranka
(1971 a. b).
9. Survival in the Preferred Habitat
‘The habitat preference of Amphibolnrus
moculosus is unique among vertebrate animals
and many factors would appear to count
against survival im such a harsh. extreme
environment, There jire obvious problems in
maintaining water balance, arising from the
hygroscopic salt surface on which the lizard
spends much of its time, compounded by lack
of aceceys to free water from which to supple-
ment its supply, and the need to handle a
higher than average sodium intake. (The ants,
Melopherus sp.. have uw high salt content
(Braysher 1972!) and many of the insects
(rapped on the lake surface huve salty encrus-
tation on their body and legs), The survival
oF an insectivorous animal in a Vegetation
free habitat, periodic flooding of the lake.
high summer lemperatures, and lack of cover
from the sun, are all matters which require
spectul physialogicul of behavioural siljust-
ment
However the liza has the ability Lo &xploil
this habjtat successfully. The low rate at
reproduction (one to four egeys per female per
anium) sugeests that its biology is more than
adequate to meet the environmental stresses.
As with all desert-adapted animals, behavioural
avoidance of environmental extremes (Schmidt-
Nielsen 1964) has been shown to be a key
factor jn survival in this harsh environment,
hut the study also indiested that the enviren-
mental stresses encountered require consider-
able physiologicul adjustment to ensure sur-
vival. The wide scape ol this study has only
permitted superficial examination of these
adaptive mechanisms. and more detailed
studies have been undertaken by other
workers,
Taleranee 1 teniperadive ane water stress
Survival of Aimphibolurus pacniosus in the
lake Eyre habilat involves ability to with-
stand high temperatures and to cope wilh the
lack of free water,
Preliminary experiments were carried oul
16 determine some of the thermal critena and
rates Of evaporative water Joss. Although the
results are based on very few reuclings, the
data are presented here since they ure the
only records available for this species, Com-
parison is made wuh published data tor two
ather species of Amphibolurus: A. pictus
Peters and A. inerinis (De Vis), Both of these
species are widely distributed in desert areas
and oecur in the sand dune country ailjacent
to Lake Eyre. A. tnermix is generally found in
red sand and A. pielus shows preference for
white sand. Comparisons between A, plac
logs, A. picmn and A inermis are summarised
in Table 2,
a
i Braysure, M. 1. (1972).—-Water and electrolyte balance im the agurmid ligated Amphibolurax
mocntosny. (Mitchel. and the structure and function of the nasal solt gland of the sleepy lizard,
Trachivdesanrus rugesus (Gray), Unpublished Ph.D-
Vhesis, University of Adelaide.
ECOLOGY OF AMPHIBOLURUS MACULOSUS él
TABLE 2
Thermal criteria and rates ef evuporative water loss for A, maculosus, A. pictus and A, inermis
A. maculosns : A. piewus ~ Aimermis -
Mitelich, Miicach, Warburg Mitchell, Warburk = Lichr- Rawson Bradshaw & Ueatwole Rradshaw
present study present study f1965h) present study 9nsh) & Shoemaker Main (1968) (1970) (1970)
T1966)
Eouritie sya”
termnerabune
CT™ aa,9°> a5" ase 485°C 44,550 465°C
Lethal 495°C 49.6°°
feMpPeralu re
Survival 300 min a B40 min 480 min. 103 min. hI+G.As
£59 ar4gzec at 44ec an46e°C —omtins at 46°C
EWL VAG meester 158 m2/2/hr O33 ns/sfhr 1.052 0.0913
ar37.sec at a7.s°C at arsec mem! hr
ar asec
The critical thermal maximum (CTM), de-
fincul by Cowles & Bogert (1944) as that
Temperuture at which a reptile loses nruscular
co-ordination. has heen determined for a oum-
her of Australian lizards (Warburg 1965 6;
Heatwale 1970).
Comparisons between these data are diffi-
cult to make because of differences in ex-
perimental procedures. Also, the usefulness of
this thermal criterion has been criticised
because some lizards: after losing muscular co-
ordination and passing into a coma, are cap-
uble of recovery if held at that temperature
or if the femperattre is lowered. Other lizards
do not recover after these muscular spasms
and die quickly, or within a few haurs-
The CTM for A. mectiosas was. determined
using only 5 specimens. "The test lizard was
placed in a beaker in an even maintained at
48°C. After 15 minutes the temperature was
raiscd at the rate of 1°C every 5 minutes,
The average CTM for A, inaculasas, derived
from these five test animals, is 45.9°C. Two uf
these (one male, one female) recovered fully
when the temperature was lowered after the
lest, The temperature from which there has
been no recovery after cessation of muscular
spasms. has been recognised as the lethal
temperature for this species.
The survival times at high temperaiures
have been revarded by some authors ys more
meaningful criteria) and these have been
determined for several Australian agamid
lizards (Warburg 1955 a, b: Licht, Dawson
& Shoemuker L966; Bradshaw & Main 19468).
Por A. mactlosus, a survival time of 6.5
hours at 45°C was recorded, and over this
period an 8% loss of body weight occured.
Texts curried out at 42°C resulted in death
between 22 and 25 hours following a loss of
6.3% of the hody weight
TF death yt this Jow level ef hody weight
loss Jef, 34-4996 Joss at death for some
iguanids (Hall 1922)} were due ta desiccation
only, it would indicate a rather low tolerance
to desiccation for A. macnlosus and its depen-
dence on retreat to the humid region above the
salt water-table of the Jake. Further evidence of
low desiccation tolerance bas been observed in
the vivattum where deaths occurred when the
artificial “water-table” was not muintained and
sub-surface humidity level dropped.
Evaporative water losses haye been deter-
mined for 2 number of Australian lizards
(Warhure 1965 a,b, 1946; Dawson ef al,
1966: Bradshaw 1970). Attempts to measure
evaporative water losses of 4. niaculosus were
carried our using test animals either fresh from
the field or which had been acclimated to a
24 hour evele involving the attainment of tem-
peratures in excess of their maximum voluntary
activity temperature. Animals were tested be-
tween 1000 and 1500 hours, their most active
part of the day.
The five animals tested yaried from 8 g ta
14.5 2 in weight (mean 10.5 g), Although the
meusured EWE, rate for A. yaculosux varied,
probubly in relation to the degree of activity
of the test animal, the average value was 1,10
me/e/hre (08-1.5 meg/erhr). Of this total
EWL, pulmonary losses contributed 0,83
me/e/hr (0.52=1.15) and 0.27 me/e/hr
(0.42-0.29) was due to cutaneous loss,
Bradshaw (1970) has found significantly
lower cutaneous and pulmonary walter loss in
the desert-adapted .4. mermis than in other
species of Amphibelurus from more lemperate
habitats, His results suggest that the improved
water economy of the desert-living specics ix
due both to reduction in the metabolic rate
and to alterations in the integument.
In Table 3 comparison is mide between
cutancous and pulmonary water [oss in A.
maculoses and A. inermis.
The Jaw propertion of total EWL attribu-
table to cutaneous loss in the present figures
62 F J. MITCHELL
for A, muculosus suggests that the integument
may be further modified against evaporative
losses.
More detailed work is required to accurately
define the critical ambient and body tempera.
tures, and rates and sites of evaporative loss,
in A. maculosux. However, available data do
TABLE 3
Rates of eittineous and pulmonary water lossy in
A. maculosus and A. inermis
A. maculosus A inermis
iT 5 6
hody Weight 10.5g 24.1+0.97¢
total RWL 1.10 mg/g/he 1.05£0.0913
mg/g/hr
cltaneous 0.27 mg/g/hr 0.452-0.0933
mg/g/hr
pulmonary 0.83 mg/g/hr 0.57=0,0778
mg/e/hr
c/P ).33 0.80
suggest that its Lemmperalure tolerance is among
the highest known for Australian lizards, and
that evaporslive Water Josses are among the
Jowest recorded,
In its natural habiuu, A, ntaculosas, like
other heliothermic reptiles, uses a series of be-
havioural postures and movements to maintain
body temperature within a preferred activity
range. This activity range of ftetnperature is
higher in the field than the eccritic or preferred
temperature selected by the Jizards in a labora-
tory temperature gradient. However, except
under some conditions imposed by the social
hierarchy, the lizards can avoid intolerable heat
levels by retreat to the damp sand of the bur-
row zone, Subservient mules, kept away from
the burrow zone by the dominant tales, are
forced to spend long hours on the salt surface
with a consequent high level of body heat.
Survival under these conditions must indicate
the existence of plrysiological capacities to
withstand high temperature and to restrict
water loss, High body temperature is tolerated
passively and water is conserved by lack of
evaporative cooling mechanisms.
10. Establishment ond Defence of Territory by
the Male,
The early emergence, in tute August, of
dominant males of the previous season 1%
usually preceded by these lizards positioning
themselves just below the salt crust where they
can more quickly respond ta imcréasing tem-
peralures. By mid-September the territories
established by these dominant males are under
challenge by the younger males which have
subsequently emerged. An old mule challenged
and displaced by a young male retreats under-
ground for the remainder of the breeding sca-
son (September to late December). By mid-
October the tertitorial situation has become
fairly stable, and remains so throughout the
breeding season.
Apart trom the juveniles (siout-vent length
<40 mim), three categories of male can be
recognised in the dominance hierarchy,
(1) Dominant: lizards which exhibit display
behaviour and fight and never retreat from
another male. These lizards develop
marked breeding coloration with bright
orange-yellow ventro-latcral murkings
grading to brilliant. reddish-orunge ven-
trally with « pale patch mid-ventrally. The
reddish-orgnge murkings extend onta the
base of the tail and under the thighs.
Sub-dominanit: lizards which exhibit dis-
play behaviour and which retreat instantly
from a dominant mule but will fight to a
decision among themselves. Among these
lizards a “peck order” is established des-
(2)
pite their individual territories. These
lizards alsn develop good — breeding
coloration.
Sufservient males: lizatds which do not
exhibit display behaviour, and) which re-
treat from all other males or rol] over
into. submissive posture on their backs if
attacked, Only very faint yellow ventre-
fateral eMlours ure developed by these
lizards.
The territorial defence of the dominant
males follows a classical pattern. Each tetri-
tory is centred upon a look-out site, usually a
amufl mound of salt 10-20 cm above the sur-
ronnding salt, or a piece of sull-encrusted urifi-
wood embedded in the lakc, The area of
territories varies about un average of 15 m
radius and is, in part, dependent on the virility
of the controlling mule. Orientation within their
own territory and that adjacent to it fs, at
least partly, by sight and the boundary between
the tennitory of two dominant males is known
ta those two males lo an accuracy of a metre
ar so. ‘The forcing of one mile over the
boundary immediately precipitates att approach
and challenge from the adjacent male, This
technique can be used to determine hierarchy
patterns in the ficld in arcas where terrilories
af dominant males are adjacent, Male TH will
ECOLOGY OF AMPHISGLURUS MACULOSUS a
flee if forced onto the territory of male I, but
will fight (and win) against male Lin bis home
territory,
The display behaviour of the dominant
males invalves a typical push-up movement in
which the forelimbs ate flexed and the whole
of the forepart of the body moves. A short
frog-like leap is followed by two quick push-
ups. In challenging another male. the gular
pouch is lowered and full threat display
follows in which the body is raised high and
compressed laterally $0 as to increase apparent
size by enlarging the profile during a lateral
confrontation (Fig. 12). This alsa displays
fully the bright ventro-lateral markings.
Usuully the mouth ts opened wide (Fig. 13).
This display is usually adequate to deter in-
truders [rom entering the territory. When male
to male fighting does occur an established pat-
terh of events is observed. There is long-range
recornitean and challenge at distances of up to
4m. then the commanding dominant closes
the distance until a counter challenge is issued
ata distance of about 3 m, after which the two
males approach each other to a distance of
about 0.5 m. There follows up to 10 minutes
of bluff behaviour, side stepping, continual
facing-up and counter-facing in an effort ta
get the tai] into position to lash the oppanent’s
head and forelimbs, and the head in a position
where it is possible to bite the opponent’s hind
legs and buck of the tail (Fig. 13). Head-on
encounters also uccur with the opponents’
jaws becoming interlocked in tenacious biting
In an cleven minute encounter, the langest
ohserved, between males T and I ia the hier
archy, three physical clashes occurred. The
biting was directed at the hind limbs and each
clash was Over in an instant, the attacker being
flung vetticully by the momentum of the
tunge and the evasive endeavour of the
opponent,
There may be overlap in the territories estab-
lished by the sub-dominant males. While the
dominaats remain “on guard” in their terri-
tory throughour most ot the day, the sub-
dominant males adjust their emergence times
so that only one ts active within the territoty
at one time.
Subservient males adjust their emergence
limes to periods when they are less likely to
be pursued by the dominant males—in the
heat of the day or late in the afternoon. Their
level of tension in the presence of the dominant
male is reflected in their respiration rate which
may be as high as 120 breaths/ minute as com-
pared with 35 breaths/minute for the domin-
aint. They spend a very limited amount of time
on the surface and then avoid recognition by
flattening themselves, with head down, against
the sult and remaining motionless for long
periods. It challenged they immediately turn
Over onto their backs in submissive posture.
Once ground temperature excecds the
threshold the dominant males will emerge at
about the same time (10 minutes) each morn-
ing independent of light intensity oF
temperulure, Heath (1962) records a similur
temperature and light-independent emergence
in Phrynosante, suggesting the presence of
endogenous circadian rhythm. During the
breeding season this emergence lime gels
carlicr by about 30 minutes each month. Aftce
the breeding season emergence becomes rcan-
dom and the dominant males generally spend
more time underground, At thes time there is
an increase in acuvity of the subservient males
which spend longer periods active on the sur-
face of the lake, There is a lowering of ten-
sion between dominants and subservients (re-
flected in the fact that respiration rates are
simitar for both), and the subservients are Jess
inclined to retreat underground or to remain
“froze and thereby inconspicuous for Tong
periods, Territorality is not actively enforced
after the end of December and, while the
dominant male spends considerable periods
underground, the subservient males embark on
an active period ef feeding which involves
frequent excursions out over the lake tn search
of the unis which, at this fime of year. con-
stitute the main food item. In the absence of
the dominants, the subservient males may
establish territories between existing territorial
areas.
The dominant males show a renewed burst
ol activity in early April, presumably feeding
intensively before retiting underground, The
dominant males and Jute developing females
extend their activity period into May, whereas
all others retire into terpor during about the
third week of April.
The dominant males and the juveniles are
the first to emerge and last to retire both
daily and annually. Under vivarium conditions,
with temperature muintained at 27°C for 12
months, the dominant males and juveniles re-
mated active throughout the 12 months. The
subservient and sub-dominant mules and the
females went into normal torpor despite the
maintenance of temperature, These lizards
emerged for 1-3 hours every 16-30 days. The
64 bE, J. MITWHELL
reason for these arousals from torper ts not
known.
In the confined conditions of the vivarium,
the dominance hierarchy of the “population”
is establishecl within about an hour. Hierar-
chicul structure can be readily determined by
observation of respiratory rutey which range
from 30 breaths/ minute in the dominant male
to 120 breaths’ minute in the subservient males.
Carpenter et al, (1970) have described the
display and aggression behaviour of three
species of Amphibolurus (A. barbatus, A.
retlewlarnys inermis and A. murtcais) aod com-
ment on the close similarity between the dis-
play patterns of these agamic lizards and the
Jeuanidae which haye been more Fully studied
(Carpenter 1967). The display action patterns
uppear to he sbecies-specific both in the
Ignanidae and the Agamidae, The display pat-
tenis of A. miaculosus haye not been fully
analysed hue the same range of postural
changes, ivolving head and forelimbs, des
cribed by Carpenter ef al. (1970) have been
observed. Brattstrom (1971) discusses the
range of postures associaied with social and
(thermoregulatory behaviour in A. farhatus.
11, Reproductive Behaviour
Vhe adult females of Astphibolurus
mutculosus do not emerge until some weeks
after the dominant males have established their
lerritories. First to emerge at 21"-23°C in mid-
Seplember are the older females, followed,
through to mid-October, by the younger
females. The females establish burrows around
the margin of the dominant male territories
and do not, at this time, move far from the
burrows. A small group of six to-eight burrows,
all within a metre or so of one another, usually
indicates the presence of a female. At this
time of year the burrows are frequently Te-
occupied and enlarged whereas later, in the
hotter weather, « burrow is seldom used twice.
During the first week or so after cmergence
the females are pot ready for mating, and
adopt two methods to repel the advances of
a male. ‘he ficst of these is circumduction, All
females of reproductive xize (>45 mm) cir-
cumduct with either forelimb in the presence
of a male, and the rate of circumduction 1s
accelerated if they are approached by the male
or come inty competition with the male for
food (Fig, 14), Secondly, should circumduc-
tion fail as a deterrent, the female twists over
on her back and lies immobile. Both these
manoeuvres serve to distract the male hy des-
troying the “female image” to which he has
responded,
Most muting activity, and much of the terri-
torial fighting, takes place at 34°-36°C, several
tlegrees below the temperature (39°C) at
which dhe body is maintained by thermerceu-
latory behaviour. Consequently during the
mating season {October to Deceniher) greatest
activity occurs between 0900 ond 1100 hours,
While feeding and terjtorial defence take
place at 39°C, preoccupation with thermo-
Tegulation prevents sexual activiry.
The male upprouch to the female usually
begins with the male elevating his head to
maximum height in order to confirm identifica-
tion of sex. He then undertakes a series of
energetic head-bobs, followed by one or two
“frog-leaps” during his rapid approach to the
fernale. A receptive female turns slowly from
the oncoming mule und wails to be overtaken.
The male approaches the female directly from
behind and with his jaws grasps her by a fold
of skin just behind the occipital region. Using
this nape grasp, the male rolls the female
over On her side. With the tail of the male
under the tail of the lemale, the cloacae lie
¢lose Logether and a hemipenis is inserted, The
pair remain rolled on their sides for the dura-
tion of copulation (about 25 seconds) with
the body of the female arched hack with fore-
limbs clear of the ground (Fig. 15}.
Folluwing copulation, the female usually
lick in a subservient position, with the head
flat on the salt, for 15-20 seconds before
moving. During this time the male “frog-leaps”
away. Females undertake weak head-nodding
during the breeding season, invalving a simple
dip and rise of the head, The exact purpose of
this is unknown bul, in view of the cryptic
coloration, it may possibly serve to muke
known their presence ta other lizatds.
Following ovulation and ferlilization, 2
number of changes occur in the appearance
and behaviour of the females. In u fertilized
fernale, ventro-lateral coloration changes fram
pearly-white to i bright orange-red, the edges
of the lower jaw become orange, and there
develop two orange patches between the fore-
limbs and two elongate orange patches along
the flanks. Also ciccumduction ceases, and
more efficient defensive behaviour ia adopted.
Upon the approach. of a male, confrontation
lakes place with the female raising the head
as high as possible and swivelling it around
to prevent the male from getting over or past
ECOLOGY OF AUPHIKOLURUS MACULOSUS 5
her If this fails, or if the male’s approach is
so fapld that this blulE is unlikely ta succeed,
the: female rolls over onto her back, frequently
well before the male makes physical contact
with fer (Fig. 16). Distracted by the changed
image the male ustially withdraws several feet,
with some head-bobbing. After 15-20 seconds
the female vighty herself, inflates the gular
pouch and stomach, wnd compresses the bady
to display the orange ventro-lateral surfaces
and so present the largest possible profile to
the inale, The female then adopts a still-legged
attitude Which hfts the body clear of the suh-
strate and with slow, deliberate steps advances
straight towards the mule. She usually passes
close in front of him, often forcing him to
withdraw a few steps or to transfer his body
back on to the hind limbs, lifting the front part
of the body to allow the femule to pass close
under his snout, After passing the male, the
female continues to walk with the stifflegped
gait, stopping on each rise in the salt crust
to look back at the male over her shoulder.
When qhout 6 m away, her pace quickens and
finally she relaxes and runs at high speed over
the salt lo disappear behind « fold of the salt
crust.
Io contrast to their timidity early mm the sea-
son, femules, once fertilized, become quite
aggressive and will attack a male should he
compete for food at close quarters. While
carrying developing eggs the female emerges at
the same time as the dominant niales and
spends maximum possible time in basking pos-
tures. Perhaps the orange ventro-lyteral
coloration of the female at this time increases
heat absorption from the substrate. Ventral
colour change in the gravid fernale has been
reported in the American lizard Crotaphytusr
collaris by Fitch (1956) and in Callisanrivs,
Copt:oseurus and Holbyookia by Clarke
(1965).
Evvs are laid 20-25 davs after fertilization,
The female digs a distinctive burrow for egg-
laying. Normally these lizards merely nose
their way under the salt crust and “swim” at a
shallow angle through the fine, dry drift sund,
down to the damp consolidated substratum.
The ¢ge-laying fernale carefully selects a site,
usually along the shore-line, of consolidated
sand dainp right to the surface. This egedaying
burrow is steeply angled (about 45°), extend-
ing down 21-25 em, with a distinct chamber
at the bottom in which the eggs are deposited.
Young females Jay only 2 eggs while older
ferales produce 3 or 4, After deposition of
the eggs the entrance to the burrow fs filled in
again, Observations, both in the ficld and in
the laboratory, Sugeest that each fernale digs
and fills several egg-laying burrows before she
finally deposits the eggs, Whether abandon-
ment of these early burrows is due to distur.
bance ov whether there is careful sclection of
sume particular set of conditions, is not known-
The salt content of the shore-line sund is 4-5%
and this, apparently, does not impede develop-
ment of the eggs. Attempts to hatch eggs
under laboratory conditions indicate that
maintenance of fairly high humidity during
egg development is important. Fenvales fre-
quently cmerge from egg-laying in poor con-
dition and highly desiccated (Fig, 17). The
ventro-lateral colouring fades from orange to
yellow to white after deposition of the eggs.
Hatching occurs after about 70 days and
the hatchlings (SV length 25-30 mm) first
appear in January and conlinue to emerge
uatil April.
12. Sex Recognition
Jn most animals showing marked sexual
dichromatizm it ts the male which is more
brightly coloured or stronely marked. On this
basis il was accepted, in early stages of this
study, that large specimens of Amphihelurus
maculosus with bright yellow-orange-red ventro-
lateral surfaces were male and that this
brilliant colouring was a key factor in the sig-
nal pattern of male to male sex recognition
in territorial behaviour. The sex of tagged
lizards was recorded on this basis of presence
or absence of ventrolateral coloration,
Recapture of marked specimens revealed
two stages of development at which colour
changes confused this simple interpretation.
Firstly, there is the change from pseudo-female
to male colour and behaviour at the time the
mate reaches the size of an adult female, In
some cases, specimens showing weak male
coloration were observed to show the usual
female responses of circumduction and tonic
immobility. On dissection, lizards of this group
(all within 43-38 mm SV length). proved. to
be male. Recapture records revealed that, up
fo S58 mm SV length, the first-year males of
A. maculasits show the yellowish ventrolateral
markings typical of the female after egg-laying.
Lizards of this size (up to adult female size)
and coloration are repeatedly identified as fe-
male by the dominant males and are driven
oul to the margins of the colony by Lhe re-
peated unwanted advances of the males, This
ob FP, 3. MITCHELL
suggests u possible dispersal mechanism within
the population,
Secondly, some lizards which had been
mitially tecorded as female, on subsequent re-
capture showed hrilhant ventro-lateral color-
ing. Allied with change of colour, there was a
change in behaviour with these lizards now
counter-challenging an approaching male. This
change from nearly-white female to brilliant
orange psevdo-male coloration and the change
in defensive behaviour and aggression follow-
ing ovulation and fertilization have been des-
eribed previously.
While sexual dichromatism exists, ubserya-
lions Auggest that coloration has little if any-
thing to do with sex recognition in A-
macntosuy, The female undergoes considerable
change in intensity of coloration Erom com-
plete luck of ventroJtateral coloration through
pale yellow to brilliant orange afler miiting,
but. is pursued by the dominant males at all
stapes in the development of this colour pat-
tern,
A first-yeal’ male. on the other hand, may
show typicul male colouring, with prominent
rusty spols oa the shoulders and weak gular
pouch stripe and yet be repetedly mistaken
for a female, Thus, first-year males, whether
they have developed mule coloration or not,
are recognised as female within the first-yeor
female size ringe of 46-58 mm.
In an effort to determine the role played by
colour in sex recognition, and to determine
whiul colour pattern might act as an innate re-
leasing mechanism in territorial display,
domimant males in the vivarium were presented
with « range of coloured models, sintulating
male colorution. None of these elicited any
response other than an investigatory lick.
The above observations suggest size lo be
the basic factor involved in sex recognition and
suggest that size judgement is particularly goud,
In relation to clevation (most look-out sites
in the habitat of the lizards arc no more than
20 em ubove the Jake surface), A. meaculosus
shows remarkubly acute vision, Observatians
of territorial challenges in the fieldt have indi-
cated that these lizards have perception of
movement and recognition of posturing at dis-
tances of 50 metres.
Both laboratory and fiel! observations indi-
cple That there is some individual recognition
between members of the line hierarchy ¢stab-:
lishvd in the vivarium and between males of
adjvining (erritories in the field population,
TABLE 4
Relationship af size fo uge in Amphiboluruas
muaculosus
Mean SV length of A.
maéulostix (mm )
Female Male
Isl year 438 54
2nd year 59 64
3rd year fl 67
4th year 62 70
This significance of size ond accuracy of
size perception would also be a u key factor in
the effecuveness of the bluff behaviour, Unless
the lizards had such an appreciation of size and
ils Significance, the act of increasing the wren
to view by enlarging the lateral profile would
not be effective asa bluff deterrent.
13. Growth Rute and Reproductive Cycle
Over the periad of study, 376 body
measurements (SV length and tail length)
were recorded for marked and Unmarked speci-
mens.
Based on field recapture of marked speci-
mens and vivarium specimens for which age
histories were known, niean SY lengths of
Amphiboluras maculosus in relation lo uge wre
shown in Table 4. The figures represent the
mean maximum measurement recorded at the
end of the active season for cach year. Mean
SV length of hatchlings is 30 mm, It seems
likely that A. maculosny dies wt the end of the
third breeding season, at 3-34 years, No older
specimens have been recovered among the
marked population at Lake Eyre,
When adult mules emerge at the beginning
of the season in September, the testes ate Fully
expanded (9x 5 mm). Smear tests show active
spermutogenesis trom September through to
mid-December and active sperm have been
found in the vas deferens in Octoher, Novem-
ber and December, In tate Wecember there is
a rapid contraction in size of the testes to
6 x 3 mm. No uctive sperm have been found
in testes or epididymes from January Lo April.
This decline in male fertility coincides with
the onset of the period of reduced activity in
the males. Between late December and curly
March the older miles spend only short
periods on the Jake surface and Jonger periods
underground, At this time, young mules move
back into the territorial areca, subservient miles
emerge for lonyer periods and they and the
fomales, exhausted after egg-laying. embark
on tong hours of feeding, There is litde terri-
tori] defence, Ihe dominant males unly weakly
ECOLOGY OF AMPFUIBOLURUS MACULOSUS
Dominant Mate
TABLE 5
Summary of reproductive cycles in Amphiholurus maculosus
First yr, male First yr. Firs: yr amt a “Adult
female UA) female «R) Female
testes
defending (with a slight lowering of the gular
pouch) a small area immediately around their
burrow site.
The older males show a more active feeding
period through mid-March and April. During
this time the testes, together with abdominal
fat bodies, expand rapidly prior to the onset
of hibernation.
Information from recapture of tagged speci-
mens, as well as laboratory observations, indi-
cates that among the first-year fernales of the
population there is a bimodal pattern of ovula-
tion,
Females hatched during January-February
ovulate in November-December. A second
series of females, hatching Jate in April, ovu-
late in February, Both these groups of females
show maturity (as evidenced by ovulation) at
10 months. Of these 10 months, at least 4
winter months (May, June, July and August)
are spent in torpor. In the earlier part of the
active breeding period (October to Decemiber)
the dominant males mate with the adult fe-
males. Towards the end of the breeding period
the 1st-year females, down to 46 mm SV
lengths, are successfully mated by the dominant
males,
Sperm retention in the female 1s indicated by
the decline of male fertility in December and
Januars inactive | _ [
a’exien Hatching Ovulation Acilve feeding
February Inactive j
March Active feecdine Hatching tee
April Active feeding it?
Testes and far 5 Hatching
huadivs expatica '
May
June . i
Hibernalion |
July | Hibernation | Hibernation | Hibernation f Hibernation
August First
emergence’ H
Sentember Establishment of |
territory,
Spermatogenesis ; : A
4 ka / Ovulation
Octaber Spermutngencsis Tdentified as Marine
Mating with sdilt feinale, Retreats Cotour
females [to edge of terri- change to
. | torial uric pseudo-male
November Sprermutogenesis
Mating with adult | Ovulation
fetnales J nt
. 7 i aune
December Soenmatoxcncsis Returns ta Mating Euetaviag
Mating with Tet territorial
year females, _ area
Contraction of Exe-Jayine
the fact that ovulation in the late-hatched fe-
males, which had been mated in December,
does not acur until February. In March these
females lay eggs which hatch in April.
The males hatched in January and February
are sub-adult (with female coloration and be-
haviour) in November-December. Table 45
summarises male and female reproductive
cycles of A. maculosus.
Acknowledgements
The author of this paper would have wished
to acknowledge many instances of assistance
and helpful discussion during this study.
Thanks are due fo a number of colleagues who
pave assistance during the field trips, and to
the late Elliott Price and the families at
Muloorina for their help and hospitality on
many occasions. The assistance of Miss Carol
Pulley with long-term laboratory observations
and records is gratefully acknowledged. Miss
Pulley’s careful organisation of the available
notes and records of this work has made the
present compilation possible. The compiler is
extremely grateful to her for this. Thanks are
due also to Mr, Michael Tyler and Dr.
Michael Smyth for helpful comments on the
manuscript.
68
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.’
Fig.
Fig.
Fig.
Fig.
13.
14,
45.
16.
17.
F.. J, MITCHELL
CAPTIONS TO FIGURES 4-17
. Amphiboaluruy maculosus, showing detail of the head with deeply sunken eye, visor-like eyelid,
and absence of visible tympanic membrane.
Margin of Lake Eyre, looking north towards Prescott Point at the tip of Sulphur Peninsula,
Instruments recording continuous air and sub-surface temperatures are set up in the burrow
zone area. The beach is backed by low white sand dunes and to the left the thin distorted
crust of the burrow zone merges into the thicker, smoother salt crust of the “wet” salt zone.
. View back along the causeway towards Prescott Point (October, 1966), Quadrats were set up
along this causeway and movements of tagged lizards were recorded in the area over several
years.
- Buckled surface of salt crust on Lake Eyre. View towards Prescott Point.
. Nesis of the ant. Melophoruy sp., are regularly spaced throngh the “wet” salt zone. The
above-surface mounds of these nests are visible here as dark spots (from their shadows) in
contrast to the white salt surface.
. A. maculosus male against the disturbed base of Mclophorus nest-mound. These mounds are
used for basking and shelter and as vantage points by the lizards.
. A maculosus, overheated by pursuil, showing panting reaction typical at raised body tempera-
tures. Note deep “lens-hood” protection for the eyes.
. Salt clods on digils of a forelimb.
- Dominant male in full threat display. Body raised and laterally compressed, gular pouch and
stomach inflated.
Dominant males fighting. Specimen in background shows compressed and raised body, gular
pouch lowered, and mouth open for biting. The tails are brought into play to whiplash the head
and forelimbs of the opponent. (Dark areas at base of tail are identification marks, )
Circumduction by A. maculosus female,
A maculosus in copulation. The male biting and holding the female by skin fold behind the
head,
A maculosus female (left) showing post-fertilization colour development on lower jaw (the
under abdomen is also bright orange) and the typical defensive position. relative to the male
on the right.
Female, after egg-laying, showing lateral skin flaps under conditions of starvation and dehydra-
lion,
ECOLOGY OF AMPHIBOLURUS MACULOSUS 69
70 F. J. MITCHELL
7
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ECOLOGY
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ECOLOGY OF AMPHIBOLURUS MACULOSUS 7s
74 FE,
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Fvaporarnve water losses of seme
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VOL. 97, PART 2
TRANSACTIONS OF THE
31 MAY, 1973
ROYAL SOCIETY
OF SOUTH AUSTRALIA
INCORPORATED
CONTENTS
Rutland, R. W. R. A note on Major Structures in the Willyama Complex -
Preiss, W. V. The Systematics of South Australian Precambrian and Cam-
brian Stromatolites. Part II - - = = = a
Maconochie, J. R. Studies on some species of Hakea (Proteaceae) - - -
Kleeman, A. W., & Milnes, A. R. Phosphorian Lavendulan from Dome Rock
Mine, South Australia - - = = & A ~
Lee, D. C. Rhodacaridae (Acari: Mesostigmata) from near Adelaide,
Australia. If Ecology - - - - - ~ - “
903/
91
127
135
139
PUBLISHED AND SOLD AT THE SOCIETY’S ROOMS
STATE LIBRARY BUILDING
NORTH TERRACE, ADELAIDE, S.A. 5000
A NOTE ON MAJOR STRUCTURES IN THE WILLYAMA COMPLEX
BY R. W. R. RUTLAND*
Summary
RUTLAND, R. W. R. (19731.- A note on major structures in the Willyama Complex. Trans. R.
SOC. S. Aust. 97(2), 77-90, 31 May, 1973.
The schistosity in the transect studied forms divergent fans about the axial planes of one major,
upright, open synform which covers the greater part of the area, and a complementary antiform in
the east. This suggests a genetic relationship between the schistosity and the major structure
although the schistosity is not strictly congruous to the synform, since bedding tends to show a
dextral relation to schistosity in both limbs.
Minor folds with approximately axial plane schistosity (Group One folds), lineations, and
schistosity-layering intersections do not show a simple relation to the major folds but exhibit a wide
variation in plunge in a NNE trending zone corresponding to the dominant attitude of the
schistosity. Group One folds tend to have northerly plunges in the east of the area in the hinge and
east limb of the Mt. Vulcan Antiform, and south-easterly plunges in the west of the area in the west
limb of the Parnell Synform. The intervening shared limb of the two structures shows variable
plunges from moderate northerly in the east through reclined to southerly. The schistosity-layering
intersections show a much wider variation in any one sub-area than the Group One folds. Group
Two folds which fold the schistosity show a closer correlation with the schistosity-layering
intersection which may therefore control their orientation.
The complexity of this pattern is apparently related to an important earlier major deformation
episode, which produced major isoclinal folds. Schistosity sub-parallel to layering in the limbs of
the major structures may belong to this earlier episode.
A NOTE ON MASOR STRUCTURES IN THE WILLYAMA COMPLEX
by R. W. R. RutTLAnp*
Summary
JkuTLAND, R. W. R, (1973).—A note on major structures in the Willyama Complex. Trans-
R. Soc. .S. Aust. 97(2), 77-90, 31 May, 1973,
The schistosity in the transect studied forms divergent fans about the axial planes of one
major, upright, open synform which covers the greater part of the area, and a complementary
antiform in the east. This suggests a genetic relationship between the schistositv and the major
structure although the schistosity is not strictly congruous to the synform. since bedding tends
to show a dextral relation to schistosity m both limbs.
Minor folds with approximately axial plane schistosity (Group One folds), lneations,
und schistosity-layering intersections do not show a simple relation to the major folds but
exhibit a wide variation in plunge in «a NNE trending zone corresponding to the dominant
altitude of the schistosily, Group One folds tend to have northerly plunges in the east of the
area in the hinge and east limb of the Mt. Vulcan Antiform, and south-easterly plunges in the
west of the area in the west limb of the Parnell Synform. The imteryening shared limb of the
lwo structures shows variable plunges from moderate northerly in the east through reclined
to southerly. The schistosity-lavering intersections show a much wider variation in any one
sub-area than the Group One folds. Group Two folds which fold the schistosity show a
closer correlation with the schistosity-layering intersection which may therefore control their
orentation.
The complexity of this pattern is apparently related to an important earlier major deforma-
tion episode which produced major isoclinal folds. Schistosity sub-parallel ta layering in the
limbs of the major structures may belong to this earlier episode,
Introduction
Early work on the major structure of the
Broken Hill District is summarised in Andrews
(1922) and King & Thomson (1953). The min-
ing, companies in the district have also carried
out much regional mapping, some of which is
incorporated in the new map (1971) of the
New South Wales Geological Survey.
The published regional syntheses referred to
above are based on mapping. carried out before
the development of modern methods of struc-
tural analysis. They also tended to rely on the
dubious assumption that certain distinctive
lithologies (especially amphibolites and gameti-
fergus gneisses) could be treated as strati-
graphic marker bands. [t is unlikely, in fact,
that rocks of probable igneous parentage such
as amphibolites. were always stratigraphically
conformable, or that genuine meta-sedimentary
or meta-valcanic units were originally persis-
tent layers. The possibility of transposition of
Jaycring, during the complex deformation his-
tory must also be recognised, Extrapolation of
rock units through the large areas of poor ex-
posure is therefore hazardous.
Nevertheless, certain broad regional differ-
ences in rock type and in metamorphic grade
are well known, and the significance of these
has been discussed recently by Vernon (1969)
and by Hobbs e al, (1968), The review by
the latter authors offers no discussion of the
tegional structure although they make some
general comments in their discussion of the
structural history of the orehody.
Habbs (1966) and Hobbs er al. (1968) dis-
tinguish between Group One folds which have
“the widespread schistosity of the area as the
axial plane schistosity and a lineation defined
by needles of sillimanite parallel to the fold
axes” and Group Two folds which “deform the
oldet axial plane schistosity and often fold the
sillimanite Tineation”. Hobbs ef al. also state
* Department af Geology and Mineralogy, University of Adelaide, Adelaide. 8. Aust. 5001.
78 R. W. R. RUTLAND
that “Throughout the entire Broken Hill region
these (Group One) fold axes and the uxsso-
ciated lineation are observed to plunge fairly
consistently towards. the SW at low angles”.
On the basis of these statements. it should be
possible by making a study of schisiosity-
Jayering relationships to establish the broad
Group One structure of the region relatively
quickly, and the present study was. made to test
this possibility. [t was recognised, however. that
there might be more than one period of schis-
tosity formation in the region and that Group
One folds im one locality might be of the sume
age us Group Two folds im another. It was
decided therefore to try to provide continuity
of data on the style and complexity of deforma-
tion for a transect right across the region. I[t
wus thought that this would permit structural
and metamorphic correlations ta be attempted
and would serve to indicate productive lines
of further research. The method used would
not be very useful if transposition were of great
importance so that bedding and schistosilty
were often parallel and facing variable.
The transect chosen for study (Figs, 1 and
2) lies between Mt, Franks and The Sisters.
aid thus runs from lower grade, amphibolite
facies, rocks in the West to granulite facies
rocks in the east (Binns 1964), In this tran-
sect there is relative continuity of outcrop, The
transect was divided into sub-areas which were
studied by Honours students working in pairs
under the writer's supervision, Study was con-
centruted on the metasediments and in parti-
cular on schistosity-layering relationships.
te FUROR META Ws
[3t3¢
——Y-—. Syaciine
| —S— Antictine
aya!
South Wales
South Aust al =
Hew
Locality map to shaw the area of the
transect studied (Fig. 2). The approxi-
mate axial traces of folds postulated by
King & Thomson (1953) are shown.
R = Mt, Robe Syncline; A = Apollyon
Anticline; S = Stirling Vale Synclme:
OW and CE — Corruga West and Cor-
ruga East Anticlines.
to SANCO LLEN
syste
"
“ADELAIDE
1 ‘
‘ yoke — od
os. , de
Be ts 7 = ast
kaa uaa Sete ane i r } } Mb,” apaths,
Loo I a feo r! gy. ily a
iw
Ut \
J \ ci : 7
" I pe 3 “jo off, le ES
v Rayteals 1 3d Herasna ony Ww %
“i ot \. Z a ii Pa Bs aii i
hots ed i d + ‘owe
Mey NV me " \
ay aa a mA Vit \ Pool baie
P. 70 ! r $? | A
Fe oN
__ SEE a =i
——. Sat a Layering
—--- Vrach ' | Sphistasity
my p
V Sub-arem numer t L r 4 ss brow Lo ciregtyer
—— Gilb-arrea Ciundary, Kn — Greup lt thd aa
—-—- Denlogisal scundary Goprar) — Gegup OD tai avis
Srructura/ Clements of @ Fruntecr of @roken Hill Asgron wetween Mount Fronaks and The Sisters
Fig. 2
the data collected in each sub-area (Fig. 3).
Structural elements of the transect. The elements shown are representative attitudes based on
MAJOR STRUCTURES IN THE WILLYAMA COMPLEX 9
Schistosity-layering intersections were men-
sired or calculated at numerous localities und
can be compared with independent measure-
ments of Group One wand Group Two fold
uxes and lineations, The work was of a recon-
Naissanee nature only but Ie has produced in-
teresting new results which have provided the
hackeround for the further work now in pro-
uress.
Structural elements
It ss believed that most of the /ayering mea-
sured in the meéta-sediments corresponds to
orginal bedding (S$). This is clear when the
lithological variation is considerable and when
the layering lies at a significant angle to the
schistosily. Often, however, the layering and
schistosity are vinlually parallel and some
transposition may have occurred,
Metamorphically produced layering is usu-
wy characterised by a regular alternation of
two, and only two, distinct litholagies. Com-
monly, for example, thin mica- or umphibole-
tich laminae are separated by thicker quarto-
feldspathic laminge. Such a metamorphic layer-
ing was sutaly observed in the meta-scdiments
and then was parallel to the main schistosity,
The main urea of Granite Gneiss cust and west
of Britannia and Scotia is also characterised by
such a layering and Jocally this layering has
become involyed in Group One as well as
Group Two folds. It is not strictly parallel to
the main schistosity and thus appears to pro-
vide evidence for a metamorphic deformation
episode eurlier than the main schistosity of the
area. It is proposed to make uw separate special
study of the granitic gneisses in the transect
studied.
Schisrasity has a remarkably similar atlitude
over the whole region, Schistosity layering in-
tersections often do not correspond with Group
One fold axes or lineations however, and It
seems probable that the deformation respon-
sible for the present schistosity attitude was
superimposed on earlier deformations, Locally
therefore the main schistosity (S,) may rep-
resent 4 but little modified carticr schistosity.
Folds of a later episode which are generally of
Group Two style alsa locally develop a new
saidl plane schistosity (S.} of similar attitude,
Thus the schistosity of the region although of
fairly constant attitude may be the composite
result of at least three deformation episodes,
(There is ulso fourth schistosity developed m
the retrograde schist zones.)
Folds can be classifird into three geometrical
types uccording to their relation ta the local
schistosity. Group 1 and Group 2 folds are as
defined by Hobbs (1966) excepi thal it is not
implied, as required by the definitian of Hobbs
ed al, (1969) that mineral Jincations are paral-
lel to Group One fold axes and indeed this is
often not the case (Rutland 1969. Anderson
1971). Folds earlier than Group One can
scurcely he recognised if the Group One schis-
josity is parallel to their axial planes but where
the schistosity has been superposed ucross their
wrxial planes, they can be distinguished as
Group Nought felds-
In any one locality the age relations of the
three groups of fulds are clear but cdetermina-
lion of aye telathonships over a larger area
depends on the correct identificution of the
associated schistasity, TE most Group One folds
are related to a single deformation episode
{D,) responsible for the main schistosity (Sy }
there may nevertheless be earher Group Onc
folds (D,) where an caclier schistosity (S,))
has been preserved and later Group One folds
{D.) where a new schistosity (S,) has locally
obliterated the main schistosily,
Major structures
The sinicture of the transect studied is do-
minated by a major Group One syntorm, here
called the Parnell Synform (Fig. 2) since it
appeat's to correspond with a fold recognised
on different grounds, and named the Parnell
Syncline by Cordwell (unpublished map for
New Cansolidated Goldfields, 1962). The axial
Irace Hes betweeli the Corruga West and Cor-
ruga East anticlines of King & Thomson
(1953), In ihe east the hinge of a Group One
antiform complementary to the Parnell Syn-
form has been recognised and is here called
the Mt. Vulcan Antiform. According to Cord-
well it represents the northern end of his Dar-
ling Range anticline (the Darling Runge Basin
of Andrews (1922)),
in the west the transect has been extended
only as far ay Mi. Franks. North and west of
Mt. Franks, Anderson (1971) hus mapped a
major structure. the Mt, Robe synform which
falds the only schistasity recognised im the
area. However, no clear evidence has been
found for a major antifarm between the west
limb of the Parnell Synform and the east limb
of the Mt. Robe synform although King &
Thomson (1953) show an anticline in this
region (Fig, 1}. the Apollyon anticline, which
they describe (p. 550): only as a “broad {l-
St) R. W. R. RUTLAND
defined anticlinal structure”. Elucidation of the
relationship between the Parmell and Mt. Rohe
synforms, which may be a faulted relationship,
must await the further work which is now
Planned, In the most westerly sub-area mapped
the lavering generally dips east across a steeper
schistosity and thus it appears 10 correspond
to the west limb of the Parnell synform.
In Fig. 4 the angles of dip of Jayering and
schistosily of individual localities are related
to cach other. It emerges that in the region be-
iween the axial traces of the Mt. Vulcan and
Parnell folds, both schistosity and layering tend
to dip west, but the schistosity fs steeper than
the layering (Fig. 4b). Across the hinge zone
of the Parnell synform both schistosity and
layering change altitude so that in the west
limb the schistosity dips custerly more steeply
than the Inyering (Fig. 4c), Similar changes
take place across the hinge zone of the ML.
Vulean antiform (Fig. 4a), The schistosity is
therefore not strictly axial plane but is a gen-
erally congruous divergent type in both unti-
forms and synforms, It is therefore clear that
this schistosity (defined as $,) was generated
during the formation of the Parnell Synform
and Mr. Vulean Antiform,
It showld be noted, however, that the obser-
vations plotled in Fig. 4 are biased in favour
of Ipcalities where a clear difference in attitude
berween layering and schistosity was observer,
In many localities, especially in the east limb
of the Parnell Synform and in sub-area I of
jhe west limb, layering and schistosity are
nearly parallel, The Parnell Syntorm, how-
ever, is a relatively open structure so that these
relations are perhaps not fully explained by
the divergence of the S, schistosity. This work
Lberefore suyvests the possibility. that the schis-
tosity parallel or nearly parallel to Jayering
may represent a but little modified earlier schis-
fosily (3,) corresponding to an earlier defor-
matian episode)
Analyses of the structure of the Broken Hill
Tegion hive suggested the presence of vurious
other major folds in the limhs of the Parnell
Synform. Some congruous subsidiary folding
has beea recognised on the limbs of the Syn-
form but in general, if major folds such as the
postulated Corruga Hast and West anticlines
exist in this transect (Fig. 1), the schistosity
layering relationships suggest that they must he
of an earlier generation (i.¢, D,) than the
Group One Parnell Synform (D,) and its as-
soctaicd S} schistosity,
For the purpose of further description the
transect has been divided into four wnits, viz:
(1) The Sisters-M1, Vulcan area: the Mt. Vul-
can Antiform
(2) The Maybell area: The Parnell Synform
(3) The Springs-Old Mc. Gipps area
(4) The Mt. Pranks area.
(i) The Sisrers-Ms. Vulcan area; the M1.
Fulcan Antiform
This area is occupied hy a group of fairly
uniforms quurtco-feldspathic xemi-schists, poor
in mica. Neither layering nor schistosity is well
developed so thin collection of structural data
is difficule, Amphibolite bands occur focally
and appear to be roughly confarmable and the
group has been intruded prier to the develop-
ment of the main schistosity by a roughly con-
cordant body of coarse granitic gneiss with
lurge feldspars.
The main antiformal hinge zone has been
recognised in the area WNW of the Sisters in
a km wide belt close to the Terrowangee un-
conformity (Sub-area IV, Fig. 2, and Fig. 5),
The antiform bere is clearly a Group One
structure with nearly vertical schistosity of
similar strike on both sides of the uxial trace,
The diagram (Fig. 3, sub-area TX) of poles
f the layering indicates 4 northerly plunge of
about 40" in general agreement with ihe trend
of miner Group One folds, The main schis-
tosity of this Group One antiform will be de-
fined as S, and the oge of the schistosity in
other localities will be discussed with reference
to it.
The Sisters Synform which is clearly out-
lined by the Quartz-magnetite rock mentber
fsee Figs, 2 and 12, King & Thomson 1953)
may be a complementary synform to the Mt.
Vulean Antiform or, more probably, an carlier
isoclinul fold on which the §, schistosity has
been superposed, The schistosity in the western
limb of The Sisters fold appears to be appro-
priate for a congruous axial plane schistosity
but the near verlicul schistosity does rot
change in altitude across the hinge of the fold,
and thus crosses both limbs (which both dip
cast} im Lhe same sense, A complicating factor
is that the schistosity must have undergone
some modification during the defermation
which produced the parallel cleavage in the
overlymy Torrowangee. but it does scem likely
1 A cleur case for two episodes of high-grade schistosity formation based on superposition has now
heen established from subsequent work in adjacent areas (Rutland er ai. in preparation).
DIP OF SCHISTOSITY DIP OF SCHISTOSITY
DIP OF SCHISTOSITY
MAJOR STRUCTURES IN THE WILLYAMA COMPLEX 81
Ro
eK
SUB-AREA Vii} 50 we *
as °°
Le EAST LIMB
* aie oa MT. VULCAN
+ So e
- 70% ANTIFORM
a Q 2" ‘a °
" reo. ‘
70 50 30 10
ee
wn
°
3
4
WEST LIMB ,
MT. VULCAN ; uA ms a
ANTIFORM #3 50
SUB-AREAS V & VII a4 /
>
s
aw
=| EAST LIMB ye FS
$| PARNELL *. ‘
SYNFORM /. b
. A 50
SUB-AREA IV \ 506 i ; WEST LIMB
ie / PARNELL
SYNFORM
Easterly
EAST LIMB
10 .
PARNELL
SYNFORM P 7 a
Westerly
Westerly Easterly
DIP OF LAYERING
Fig. 4. Relative dips of layering and schistosity.
= RK. W. KR. RUTLAND
that the Sisters fold is of an earlier generation
than the Group One Mr. Vulcan antifurm and
the associated S, schistosity, and there is clear
evidence of minor folding earlier than the
schistosity..
Most minor folds in the hinge of The Sisters
synform are Group One structures of distinctly
Variable mortherly plunge, One excellent ex-
ample of earlier minor folding occurs on ihe
hinge of u Group One structure, The earlier
fold on which the schistosity has been super-
posed has a nearly horizental plunge and an
east-north-east trend. Conceivably therefore
the main Sisters fold is a reclined carlicr struc
ture on which the mami schistustty and usso-
ciated Group One minor folds have been
superposed. Vernon (196%, p. 52) also notes
that “small isolated jsoclinal fold hinges in
quartzite fxyers can he traced around the
hinges of the relatively large folds” at the Sis-
ters.
Further south in the traverse ESE of the
Springs H.S. (sub-area VIIL, Fig, 2) a Further
complication occurs with the recognition of
two schistosities. The muin schistosity generally
dips north-west in the western limb of the Mc.
Vulcan antiform. In the north draining valley
about two miles east of the Springs H.S., how-
evec, both the layering and this schistosity are
folded round the hinge of an antiform whieh is
therefore Group Two with reference to this
schistosity, A new schistosity is locally deve-
foped in the Hinge, however, so that with
reference to the later schistesity the fold is
Group One.
Ik is not clear which of the sehstosines
should he correlated with the single prominent
schistositv immediately to the north. If the
earlier schistosity is so correlated (as is fay-
Oured by its intensity) then it may be described
as 8, and the Jater schistosity can be regarded
asa loeul development of S.. Lf, however, the
Jater schistosity is correlated with that to the
north (as is favoured by their orientatians)
then the earlier schistosity must be regarded
as one preserved from an euslier period of de-
formation, ic, it would be an S,, schistosity.
Purther work is needed to determine unequi-
yocally whether the major antiformal structure
in refiuled to the earlier or the Inter schistosiry
but the seoond alternative is here preferred,
Further south again in the traverse between
Razer Back and Moorkaie the schistosity-
luyertny relations indicate the presence of o
Grup One synformal structure east of the
fai uniform (Fig. 2), West of the Springs
H.S., however, layering again dips west more
gently than schistosily and the Moorkaje syn-
form is therefore regarded ag a relatively xub-
sidiary fold on the west limb of the major Mt.
Vulcan antiform.
It is notable that both Group One and
Group Two folds and lineations in this region
have northerly plunges (Fig, 3, swb-areas WIT.
VITT and EX). Schistosityayering intersections,
however. are much more vuriable and may
have any plunge within the NE striking zone
corresponding to the general allitude of the
schistosity.
Tt is inferred, therefore, that the Mt. Vulcan
antiform as defined, is a Group One structure
superposed On tsoclinal folding represented by
the Sisters Synform. It is evident from the
relations desenbed, however, that this Group
One structure could correspond im aye to
Group Two structures elsewhere, which fold
an carlier schistosity. It cannot be assumed that
the antiform is an anticline or that it folds a
simple succession of constant Facing,
(2) The Maybell areat the Parnell Synform
The hinge zone of this structure occurs in
sub-area IV in a group of mica schists with
quartzite and quarty-rich gneiss interbeds, Both
schists und gneisses are commonly garnet-
ferwus.
The well-developed lavering and schistosity
in these rocks has facililated folding which is
tighter and of smaller wavelength than in the
Mt. Vulcan antiform. The hinge zone is made
up of several folds so that the main axial trace
is not easily drawn without detailed mapping
(Fig. 6). The map suggests that the axial trace
has suffered some right-lateral offser on the
ENE trending shear zones which cross the area,
The diagram of mesoscopic data for this
structute (Fig, 3, sub-urea TV) is somewhat
diffuse so that a plunge variation from gentle
north to gente south can be inferred, At least
part of the spread is due to Group Two folds
aml these generally plunge to the south. The
main fold, however. is clearly a Group One
structure and minor Group One folds more
commonly plunge to the north (Fig. 3). Bvi-
dence along the main hinge line in the northern
part of the area for example gives a plunge of
16/020 while dextrul folds on the west side
of the gneiss in the south of the area plunge
$4°/028. A few hundred yards west of the
latter outcrop, however. Group One folds in
interbedded qnartzites and schists plunge 30"/
196, Mineral lineations have occasionally beca
observed to plunge north at moderate pngles
83
MAJOR STRUCTURES IN THE WILLYAMA COMPLEX
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in both limbs of the major structure but others
have been obseryed to plunge gently to
moderately south.
The Jayering-schistosity intersections again
show much greater variation than the fold axes
(Fig. 3). This shows that strain axes and peo-
Metric symmelry axes were not coincident and
suggests that the schistosity may be superpased
on i complex and possibly pelvphase fold sys-
lem, The clearest evidence of this ts in the pre-
sence, in the gneisses, of what appears to be u
melamorphically produced Javering which has
been folded with the main schistosity as uxial
Plane. Tf the main schistosity 1s again defined
us Sy) und the associated Group One folds as
D, then the metamorphic layering provides
evidence of an earlier Dj, metamorphic ude-
formation episode:
In sub-earca IW (Fig, 2 and Fig. 3) schistosity
and layering trends are roughly parallel,
though dextrs) relations are more commen in
the west limb amd sinistral relations in the east
limb, suugesting « closure to the nurth*. Over
a wider area, however, 4 notable feature is that
the liyering tends to show w dextral relation
to the schistosity in. both limbs of the fold, ic.
in sub-areas ILL und Vo (Fig. 2). ‘This would
imply a generally southerly plunge of the schis-
tosity layering intersections in the west limb
and a noltherly plunge in the east limb. In
fact Croup One fold axes do commonly have
a vorherty plunge in the east limb althoug’y in
sub-alreas Yoaod VI the plunges are often very
steep, Group Ong folds in the west linib (suh-
wea VW) da have a. south-easterly plunge It
can be inferred therefore that the schistosity
lies slightly incongruously across the structure.
though Lhe divergent attitude ef ihe schis-
tosity with respect to the axial plane confirms
That the major structure and the schistosily are
Beneticully rehited,
The limb areas of the Parnell Synform ure
Jiscusseed tn more deta! below
(A) Pie Sprentge-Oled Mt. Gipps arent
Schistosity and layering are offen nearly
purailel in this area bute where their dips have
heen distinguished the layering generally dips
west less steeply than the schistosity, The schis-
losity-luyering intersections show the usual
vanation in i NE trending great cirele, In sub-
urea V (Fig, 3) the schistosity-layvering inter-
sections more commonly plunve steeply ta the
south us do observed mineral lineations in the
schists. Group One fold axes, however, vury
from steep south-eusterly plunges to ‘steep and
moderate northerly plunges while Group Two
folus vary from steep southerly to stecp mor-
therly plunges. On the eust side of the main
franite-gneiss body, however, in the schists of
sub-aren VIL (Fig. 3) schistosity-layering inter-
sections plunge less steeply cither NNE. or SSW
and observed Group One fold axes have exclu-
sively NNE plunges. This contrast between the
variable plunge of schistogity layering inter-
seclions and the mofe consistent plunges of
Group One fold (xcs is similar to thal already
neled for the hinge area of the Purnell swoforin
abd reinforces the suggestion that the main
schistosity is slightly oblique om the major
simucture.
In the cast limb of the Parnell Synform no
euther major folds (Group Nought) have heen
recognised hut if the folds postulated by King
& Thomson (1953) and Cordwell (unpublished
mup for New Consolidated) Goldfields. 1962)
cxist they would belong in this category, Brief
coniments. On these postulated folds ure mace
helow,
(a) Tue Muiaas Syncrine,
This fold was inferred to lie with uw NNE
trending axial trace in the schists in sub-are:
VIL (Fig. 2) east of the miin body of eranite
gneiss. There is no evidence of repetition of
the sticcession in the meta-sediments on either
side of this trace although the acromugnetic
map <loes sugpest the possible repetition by
folding of the distinctive quartz-magnetite rock
unit. Boudics of granite gneiss da flank the
schisis un both sides of the proposed fold bul
this rock type is probably intrusive, although
it was emplaced before the development of the
muin schistosity, Layering ind seistusity are
oflen nearly parallel in this area and schis-
tosity-layering intersections may plunge either
north or south, Where distinguished, however,
the layering dips west more vently than the
schistosity. Thus the area of this postulated fold
lies wholly in the western limb of the main Mt
“Phe sehiitosity-layering relationship is described as dextyal when the layering trace in pralite tor
on a horvzontal surface! fs clockwise fran» the schistosity wace. This any minor tuck or “drne”
folita toowhich the schistosity is wsial plane are also deaxtral and have a dexeral schistosity-livering
relalionshife ins their long limbs. Conversely the schistosity-layering relationship is described as
sinisual when the Jayerme trace In prohle for on 4 horizontal surface) is anticlockwise frou the
schistosity trae.
MAIGR STRUCPURES IN ‘THE WILEYAMA COMPLEX RS
Vulcan antiform ane, if it exists, it is earlier
that the axial-plane schistosity of hat Fold.
(tb) Yanco ANTICIINE
The main outerop of granite gneisses north
of Britannia and Scolia was inferred co lie in
the core of an anticline. Southwiirds this Yanco
anticline was supposed to be divided into two
subsidiary anticlines (occupied hy the two
southerly projections Of granite gneiss), sepa-
rated by a syneline of metusediments south of
Britannia and Scotia mine. Insufficient data has
been wathered to evaluate this hypothesis fully
but several relevant poinis can be made
(Fig. 7).
The granite gneisses display a complex inter-
layering with the metusediments, w relationship
which may be primary or due to subsequent
deformution, There is evidence of three defor-
mation episodes in the gneisses. The main
schistosily S,; (D, deformation) is superposed
on ao earlier metamorphi¢ layering in the
eneiss (D,,) and it is tolded into Group Two
folds (Do). The prominent southerly Jineauion
may be related to D, or D,.
A Group Two fold of more than one hun-
dred metres wavelength occurs in granite
gneiss ahour 1500 m south of old Mt. Gipps.
This is a large sinistral parasitic fold with a
soulherly plunge on a general westerly dip.
Other Group Two folds are dextral with oor-
therly plunges but all are parasitic folds and
there Is no evidence of any major Group Two
fold which might repeat the succession.
Schistosity-layering relationships are com-
plex: venerally schistosity and layering are
nearly parallel aod the attitude of their inter-
sections is very yuriuble: where the ungle he-
tween schistosity and layeting tx substantial the
achistosity generally has the steeper westerly
dip but in some cases the layering dips casi-
watds less steeply than the schistosity, This
suggesis The presence af sume Group One falds
and a chear example. dextral with northerly
plimge, cin be seen nearly 3 km NE of
Old Mt. Gipps. Again, however, no systemutic
vanation in schistasity-layenng relations was
found which might indicate a majur Group
One fold and in general the layering dips west
more gently than schistosity on both sides of
the main granite-gnems outcrop, It is therefore
inferred that. if the eranite-gnciss areas do tie
mm the hinges af major folds, such folds must
be earlier than the Parnell Synform and Mt,
Vulcan antiform, No positive evidence for such
folds bas been found bur Curther study of vhis
aréa is required
(7) NINE-Mi.e SYNCLINE AND MAYREDI
ANTICLINE
These fulds were inferred by Gordwetl in
the essentially metasedimentary aren west wf
Old Mi. Gipps (the Maybell anticline carres-
ponds approximately to the Corrusu Rast unti-
cline of King & Thomson 1953). They were
apparently based partly on the suppased repe-
tition of amphibolites and of a para-gneiss unit.
These repetitions have not heen substantiated
hy the present work. Distinctive lithologies af
pale calcareous schists occur along the old
tramway north-wesl of Old Mt. Gipps and
these have not been found further west.
However, schistosity and layering shaw a
variety of relationships although in general the
layering dips west more gently than schistosity
ina dextral relationship and phinges are nor-
therly (Fig, 7). Along the old tramway, haw-
ever, the layering dips east more gently than
the steep schistosity and the prominent Grougr
One folds ate dexttul on a southerly plunge (ys
in the west limb of the main Parnell syntorm),
There is some inconclusive evidence in these
outerops that the Group One folds refold un
earlier set of Group Nought folds. These rely-
tions of Group One structures surgpest the pre-
sence of uw minor dependent antifarm in the
cast limb of the main Parnell Synform, but, us
with the major structure, the plunge yarition
suggests that the schistesity is oblique to the
axial plane of the fold.
(4) THe Mi, Franxs Ages
Tn sub-areas Wf and 1 the layering generally
dips east more vently than the schistosity in a
dextral relationship an a southerly plunge,
often steep. This is especially well seen in a
coarse schist unit defining a number of cextral
folds in the ares south of Florida Mine. There
is sone evidence that this anil closes sourh-
wards in a major fold wih a core wt Parnell
gneiss (resembling Potosi gneiss) but beth
limbs dip cast across the yertieul schistosity und
the Group One folds have the same dextral
sense in both limbs. This suygests that the pos-
sible major fokl is a Group Nought fold, with
reference to the axial plane schistasity of the
Parnell svnfarm, No evidence for the sorth-
ward continuation of the Sdrling Vale svncline
(King & Thomson 1953) was found in this
sibearca.
Tn sub-area Il the schistosity und Joyering
are generally nearly parallel and Group One
Minor folds are rare, Most commonly the
Inyering dips cast acrots f steeper schistosity in
86 R. W. R. RUTLAND
Schistosity - Layering
hinge zone of the
60 After
55
>
&
Fig. 6.
a de¢xtral relationship on a southerly plunge
but near the eastern margin of the sub-area
_some sinistral minor folds occur, also on a
southerly plunge since the layering here dips
east more steeply than the. schistosily. This sug-
gests the presence of an overturned south
plunging antiform-synform pair on the western
limb of the main Parnell synform. The axial
trace of the Apollyon anticline as mapped by
relationship
Parnell
in the
Synform
L.Chenoweth and L,Schmidt
i
f
A283 751
7
63 2
eo
i A co
a he 8 ay. ht
V5 BS
vss ag
—- Layering
Lj} Sehistosity
4 8B wn »=ORetrograde faylt zones
a a Approximate trace of
Parnell Synform
Vabtty o ri
L 11 1 i |
ho OM bites
Structural elements in the Maybell area: the Parnell Synform.
King & Thomson (1953) runs through this
area and further work is required.
Also in this sub-area, close to the track cast
of Mt. Franks the quartzo-feldspathic sedi-
ments show a strong “elongation” Jineation
plunging south-east in relatively low angle
schistosity. These rocks are also inyaded by
abundant pegmatite and they provide a strong
structural contrast with the tocks further west,
MAJOR STRUCTURES IN ‘THE WILLYAMA COMPLEX
Possibly the lineation and the schistosity which
is sensibly parallel to layering are preseryed
from a deformation episode earlicr than the
Parnell synform.
In sub-area I there is relatively little peg-
matite and the rocks are commonly phylilitic,
though some members are rich in andalusite
porphyroblasts, around which the schistosity is
deflected, The layering again generally dips
east more genlly than schistosity but in con-
trast to sub-areas IL and IIL the relationship
is now generally sinistrul and the plunge of
the Group One minor folds is to the north,
The Jayering actually trends ubout N-S and the
schistosity about 030°.
Thus the relationships are still consistent
with a position in the western limb of the main
Parnell synform but the trend of the layering
$7
hus changed and the plunge has changed from
southerly to northerly. These changes might be
attributed to the presence of a southward clos-
ing Group Nought fold between sub-areas I
and Jl. No evidence of such a fold has been
found and the strong contrast of structural
style between the two sub-areas rather suggests
the presence of a thrust fault.
Nearer to Mt. Franks there is a reversion
to a dextral relation between schistosity and
layermg and to a southerly plunge since the
layering continues to dip east across a nearly
vertical schistosity,
On ML. Franks itself there is a Jarge dextral
fold and crenulation cleavuge is strongly deve-
loped. It is apparent, howeyer, that the S;
schistosity which has suffered crenulation was
itself formed at a high angle to the layering.
=
N
4 _ -
IEE aT (Rad 8 EG
/, bS
/ } i Ff of ( 56 el of of! x t E | y ‘ f f
AVR Nae HONKY
NE LF INE PI ON TS 7
boa "IC Xe / wa ral la Wt (2 Me 4 \ WY , F| irs Z
io ‘ qe 4 oy Near +9 \ \ iF f af 5 / F {
a “ is ; 7 !
# \Wese AG Ps Te ) a Tee ps 2} Seat Ph - f
ae ft Op / “& Se ot wf i \ { fs
a i / LS a ein ut cies Hel f of oe
a leg AN. a ( 4) ) / f / i ah
oe , y 5 j - Bh
as} / 2
Pat AWS ¥ A i (3
f Ved a re - des
i A lp
aol FY She
os, wy \ c a toe é > ae ‘iad
by c ~ ¢ Le ¥y,
_ a ri we/ t \ fa. aa yf
~~ ey, YA to f
§ Ma ge ol af \ as {
/ = Sei)? \Y 7
ee) ‘
Pres ¥
ye |
fe so 1 }
Metres Milometees
pre Broken Hill |
=— Road >— «Group I told axis “+ Intersertion of layernny and schistosity
oe Layering > #$Groun DT told ents Rh Creriglathen cleavage
LI Sriratosiry —+ ~=Mineral broeatien ta} Areas af sa jd poet water
Fig. 7. Structural eleinents in the Old Mt. Gipps area.
8s R.
Ft is inferred therefore that the main Mt,
Franks: structure is essentially a large Group
One dextral fold modified by Group Two folds
and crenulation cleavage, As noted above, the
elucidation of the relationship. of these obser:
vations io the interpretation by Anderson
(1971) of the Mt. Robe area must awail
further work.
Conclusions
minor folds of Group Nought, Group One
and Group Two styles have been recognised
and would suggest the presence of three
periods of deformation if it is assumed that the
schistosity is everywhere of the same age. Some
doubt is cast on this assumption by the fact
that in some areas (particularly the hinge zones
of the major structures) the schistosity is con-
sistently steeper than layering and mukes a chs-
tinct angle with it while in others (especially
in the limbs of the major structures) it 1s
nearly parallel to the layering.
The Parnell Synform and Mt. Vulcan anti-
form are apparently of Group One character
since ihe undoubted $, schistosity is roughly
congrucnt and forms divergent or reversed
fans. Since the schistusity attitude varies sys-
tematically according to its position in these
folds tt seems probable that the folds and the
divergent schistosity, non-parallel to layering,
were formed during the same deformation epi-
sode, Nevertheless the fact that the layering
generally shows « dextral relation to. schistosity
in both limbs of the Parnell Synform suggests
that the schistosity has been superposed
stighly obliquely on the fold, after the initia-
tion of buckling. This oblique superposition
may be partly responsible for the difference in
plunges in the two limbs of the Parnell Syn-
form hut there is some evidence that earlier
folding is also significant.
The dominant dextral relation of layering
to schistosity sugycsts that Group One folds
related to the schistosity should plunge south-
cust in the West limb and north in the -east
limb of the Parnell Synform, In fact
schistosity-layering mtersections and fold axes
(Figs, 3 and 8) show a wide variation in att-
tude in a great circle zone striking NNE. This
suggesis that the laycring attitude on which the
schistosity was.superimposed was variable, pos-
sibly due to earlier folding. Moreover, there is
not a close correlation between the attitude of
Ww. R. RUTLAND
' ——~
4 | eh
' a
' . “a ,"
. '
“4 nf
‘ 4
/ a» ah Eee
/ Ao ? «1 7
/ os Mn \,
/ 4 ‘ =
/ . atte , ‘
al
{ .
ate
| . my \
| . Jt ¥ |
.
i a
| = ee {
. .
i} “ 4
°
\ “ap a4 y
' * Iai 2 Fs
i .* 3 ree
“4 5 . ‘
- «’ 4 . ‘ “ 2 \
. a . 4 - 7
oy . 2
- 4, , ¢
+ % ‘
4 *s
.
le 7 a Pe The
« Cevp tT tntan
Timeations
Fig. $ Lower hemisphere equal-area projection
of fold axes and lineations for the whole
Irunsect.
Group One folds and schistosity-laycriny inter-
sections, There ts a closer correlation between
layering-schistosity intersections and Group
Two folds suggesting that the latter have been
controlled by the former.
Minor folds of Group Nought character
which can unequivocally he said to be earlier
than the schistosity are rare however. The di-
vergen| schistosity inevitably approximates to
the axial plune of any eurlicr pearly isoclinal
foluls and the distinction can be made only in
areas of favourable lithology and exposure’,
I. has giready been noted, however, that un
carlicr schistosity may be preserved, and the
evidence in gneisses of u metumarphic layering
tarlicr than the S, schistosity provides con-
vincing evidence of an eurlice deformation epi-
sode. On the map-scale moreover, the Sisters
Yok and some others appear to be isoclinal
major structures earlier than the Parnell syn-
form and its related schistosity. Since the axial
planes of these isoclinal folds are approxi-
mately parallel to the limbs of the Parnell syn-
form it follows that they were recumbent
before the deformation which produced the
Parnell synform (Fig, 9),
%A good example outside the area under discussion can he seen south-east of the main road, about
onc km north-east of the Flying Doctor base, There the schistosily-layering Telationsbip is sinisuwal
and congruent Group One minor folds oceur with south-westerly plunges o! about 50". The schis-
tosity culs across occasional earlier dextral folds which plunge north-north-east.
MAJOR STRUCTURES IN THE WILLYAMA COMPLEX ay
cf PARNELL
SYNFORM
‘hrust oof
deformation
Possitie
PAbliat
Sehistosity (Sy )
cf MT. VULCAN
ANTIFORM
Axial trace of probable
wcarlier Tatds and attitude
Of #ariter sohistosity
Layering | S7
Fig 9.
Diagram to show general form of the Parnell Synform and Mt. Vulcan Antiform and their
possible relationship ta major structures of the inferred earlier generation. No attempt is
made to show specific early structures which have been postulated.
Three major deformation episodes are there-
fore inferred:
1, Dy; Formation of tight Group Nought
folds (apparently Group One if the later
S, schistosity 1s roughly parallel to the axial
pline or if 5, is preserved) including the
Sisters synform, Original attitude possibly
recumbent but now upright with steep
plunges,
D,: Development of the main folds, the
Parnell synform and Mt, Vulcan anitiform
together with their associated (S,) schis-
tosity.
3, Dy: Formation of relatively minor Group
Two folds, plunging south except in hinge
urea of Mt. Vulcan antiform.
Further work is now in progress on a fur-
ther east-west transect, immediately north of
Broken Hill. ft is hoped to test the conclusions
presented here, to extend knowledge of the
Major structures ind to correlate the deforma-
tion phases recognised with those established
re
by other workers (Hobbs 1966; Williams
1967; Anderson 1971).
Acknowledgements
The mapping, on which this paper is based.
was carried out by a group of Honours s{u-
dents of the School of Geology, University of
Adelaide in May, 1969, under the writer's
supervision. The students were Messrs. K. F.
Bampton, T. H. Bell, L. M. Chenoweth, D. A,
Couzner, T. M. Clifton, N. J. Crase, R, A,
rears, D. N. Harley, P. G. Haslett, A, N,
Larking, H. Mastins, R. K_ Netzel, P. R.
Pierce, L. E. Poole, B, V, L. Rees, P. W,
Rooney, B. L. Schmidt, R. E. Williams, A
preliminary version of this paper was circulated
as Rescarch Report No. 1 of the Centre for
Precambrian Research, University of Adelatde,
1969.
Financial assistance from the Broken Hill
Mining Managers Association is also gratefully
acknowledged.
References
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THE SYSTEMATICS OF SOUTH AUSTRALIAN PRECAMBRIAN AND
CAMBRIAN STROMATOLITES. PART II
BY W. V. PREISsS*
Summary
PREISS, W. V. (1973).- The systematics of South Australian Precambrian and Cambrian
Stromatolites. Part II. Trans. R. Soc. S. Aust. 97(2), 91-125, 31 May, 1973.
Five new forms of stromatolites from South Australia (nzeria conjuncta, I. multiplex, Jurusania
burrensis, Katavia costata and Kulparia kulparensis) are described. South Australian occurrences
of Conophyton garganicum garganicum, Gymnosolen cf. ramsayi and Inzeria cf. tjomusi,
previously known from the USSR and elsewhere, are also discussed.
THE SYSTEMATICS OF SOUTH AUSTRALIAN PRECAMBRIAN AND
CAMBRIAN STROMATOLITES. PART II
by W. V. Preiss*
Summary
Premss, W. V. (1974).—The systematics of South Australian Precambrian and Cambrian
Stromatolites. Parl If, Trans. R, Sac, S. Aust. 97(2), 91-125, 31 May. 1973.
Five new forins of stromatolites from South Australia Umnzeria conjuncta, I. multiplex,
Jurusania burrensis, Kaiavia costatu and Kylparia kulparensis) are described. South Australian
cccurrences of Conophyton garganicum gurganieum, Gymnosolen cf. ramsayi and Inzeriw cf.
tjomusi, previously known from the USSR and elsewhere, ure also discussed.
Introduction
“This paper is a continuation of Preiss (1972)
in which the principles of stromatolite classifi-
cation were outlined and several new forms of
stromatolites were described, The glossary ap-
pended fo Part I also applics to this paper.
Systematics
Group CONOPHYTON Masiov
Conophyton Maslov 1937. 334, Koralyuk
1963: pl. 5, Fig. 3. Komar, Raaben & Semi-
khatov 1965; 27, Kamar 1966: 72, Cloud &
Semikhatov 1969: 1037. Bertrand 1968: 170.
Walter 1972: 102.
Type Form: Conophyton lirietn Maslov,
from the Dereynin Suite, Lower Tunguska
River,
Diagnosis: Non-branching or extremely rarely
branching columnar slromatolites with conical
laminae, usually thickened und/ear contorted in
their erestal parts,
Content: C, cylindricunt Maslov; C. pietu-
lunt Kirichenko; C. cireulum Korolyuk:
C. garganicum Korolyuk; C. mileradoyict
Raaben; C. Jitu Maslov; C. baculum
Kirichenko; C. gaithitza Krylov; C. ressoti
Menchikolf; C. cadilnicus Koralyuk and
C, confertum Semikhatov.
Conophyton varganicum garganicum Korolyuk
(emend.)
FIGS. 1. 2a. 9a, 13, 12a
Conophyron cf. garganicus (partim),
Glaessner, Preiss & Walter 1969: 1056,
Material: Eleven specimens from Paratoo,
S. Aust.
Description
Mode of Occurrence: These stromatolites have
been found only in a diapiric raft in the Para-
too Diapir. The basal portion consists of flat-
laniinated stromatolite, passing up into large
domal structures up to 1 m diam. (Fig. 11¢).
Domes are usually laterally linked, occasion-
ally separated by small interspaces, then divid-
ing into discrete columns, 1540 cm in diam,
with conical laminae, Transverse sections of
columns round to oval or lanceolate (Pig.
Jib). Columns 1-4 cm apart, with some mas-
sive bridges, often slightly bent, with axes non-
parallel, diverging at up to 30° (Fig. lla).
Some of this divergence may be due to tec-
tonic disturbances. The original mode of occur-
rence is not clear because of the discontinuous
outcrop, it may have been a bioberm or thick
biostrome, perhaps. 30 m thick. The only evi-
dence ag to the facing of the bed is the upward
passage from flat-laminated to conical stroma-
tolites, with apices growing upwards,
Column Shape: Ficld observation shows that
columns are somewhat irregular cylinders, with
ragged edges, massive bridges and overhanging
laminae. Only one specimen was suitable for
reconstruction (Fig. 2a). Columns of round
transverse section haye a linear crestal zone,
while those of elliptical and lanceolate sections
have crestal plunes in the long axis of the
ellipse (Fig. 11b), Specimens studied in the
laboratory also show both types.
* Geological Survey, South Australian Department of Mines, Box 38, Rundle Street P.O., Adelaide
5000.
92 W, V. PREISS
‘The margin structure is very irregular. with
numerous large bumps, overhanging peaks and
shore cornices. (Pigs, 2a, lta). Bridges vary jn
thickness from one Or two to several lens of
laminae,
Branching: No true branching except actual
separation of columns from the domed and
flat-laminated base, Rarely a small projection
with convex, non conical laminae occurs. on
the margin of a column,
Lamina Shape: In longitudinal axial sections
laminae steeply conical, apical angle generally
acute (50-90°) but obtuse angles cecur near
the base of the columns, Away from crestal
zone, laminae usually straight and parallel in
longitudinal section, but in places bent down-
wards near the column margins, producing a
shupe resembling gnthic arches (Fig. 94),
Crestal Zone: All laminae more or Jess thick-
ened in crestal zone. Some light laminae
greatly thickened. Dark laminae arched up and
contorted, often leaving irregular voids filled
with sparry dolomite, within the thickened
light laminae (Fig, 11f), The erestal line, join-
ing apices of successive conicul laminae, is
very wavy, with frequent sharp displacements
of crests (Fig. aj, The overall shape of crestal
zone ts however straight (Fig. 12a); ir corres-
ponds mostly to Type HI (after Komar,
Raahen & Semikhatovy 1965, p. 23, Fig. 5)
with uneven thickenings and sharp lateral dis-
placements, but some examples of Type Il
(without lateral displacements) occur. In
places, laminae are deflexed immediately out-
side the crestal zone (Fig. !2h). The diameter
of the erestal zone is ken os the width be-
tween the limits of thickening of laminae, Out
of 33 measurements, 63% Jie between 7 and
9mm, 24% between 5 and 7 mm, and 12%
betwecn 9 and 12 mm.
Lamination; Very distinctly banded and
striated in better preserved specimens, consist-
ing of straight, parallel, smooth, very thin
laminae, either very continueus, or formed by
chains of elongated lenses, aligned in definite
layers (Figs. Tle, 11f), Two types of primary
laminae occur: light { L,} and dark (L.). In
some specimens 1. layers grouped inte fairly
Uistinct macrolaminae, in Which light laminae
are thin and subordinate, separated by layers
of predominantly L,; iype (Fig. 11f), The ap-
pearance of macrolaminae has been eNxag-
weraled by the preferential recrystallizatinn of
light laminae. L, taminae relatively pure and
transparent, mostly 0.08—-0.1 mm thick, gen-
erally of very constant thickness from the else
of the crestal zone to the column margin, never
lensing out. They are internally homogenous.
composed of xenolupic. almost equigranular
dolomite, of grain size from O.1H—-0,03 mm.
Many grains slightly incquidimensional. Occa-
sional lenticular spay-filled cavities have dark
laminae draped arvund them Fig. lle), L.
laminae darker, much fess transparent and
samewhal finer grained than L, laminae, the
Fine crystals stained by 3 pale brownish. pos-
sibly organic coloration (Fig. 1le). Mast dark
laminae 0.02-0,10 mm thick, ont os continuous
as L, laminae, frequently splitting into series
of lenses, 0.2-1.0 mm Tong, and 0.1—1.0 mm
apart, aligned parallel and separated hy pale
laminae. Some dark laminae ure continuous for
several cin; some have slight, rounded, lenticu-
jar swellings, These, as well as the lenses, may
be blunt ended, rounded, or pointed, Rarcly,
they contain significant swellings, the under-
fying ynd overlying jaminae heing draped
around them. Relatively large (0.52.0 mm)
neduies, within.a pale lamina (eg Fig. Ie)
are probably detrital carbonate grains, 1... Ia-
minac composed of equidimensional. equigra-
ular, Xenntopic dolomite, of grain size 0,006—
0.015 mm. Boundartes of L, and L, laminae
distinct and smooth, but slight recrystallization
has made them a little diffuse in plages. Mac-
rolaminac, consisting of sets of Ly laminae.
very prominent in some specimens, are U.4—-
1th mm thick, composed of 5-10 Ly-L. Ia-
mination pairs, bounded by predominantly
light macrolaminae (.2-0.5 mm thick. often
spurry and recrystallized (Fig. 11f),
Statistical Study; Numerous measurements
were made on six large thin sections, of the
following parameters: (1) thickness of light
laminae I,; (2) thickness of dark laminae L..;
(3) ratio of thicknesses of adjacent dark and
light laminae La/L, and (4) coeficient of
thickening. i.e ratio of thickness of a lamina
in crestal zone, to thickness of same lamina
outside crestal zone.
The distribution of thicknesses ef Jarninae
L, and L., were plotted graphically for thick-
ness Intervals of 0.02 oun; the [requencies of
intervals were plotted against the mid-point of
each interval, for the six specimens (Fig. 1b
to g). A comparison of the six graphs for each
lamination type shows some variation between
specimons, especially for L. laminac, which is
interpreted as being due to the difficulty of
distinguishing single datk laminae and the
thinner macrolaminge in same specimens, This
difficuity is increased with rreater recrystalliza-
lion, so that one would expect the more re-
SOUTH AUSTRALIAN
STROMATOLITES IL 93
Fig.
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1. Diagnostic characters of Canophytan: (a)—The traces of the crestal lines of two specimens
(S214 at left and $532 at right) drawn from thin section (x1/3); {b) to (g)—Frequency
distributions of thicknesses of light Jaminae Li and dark laminae Le» for 6 separate speci-
mens; (h)—Pooled frequency distributions of lamina thicknesses for all six specimens, com-
pared with data for Russian conophytons; (i)—Frequency distribution of the ratio of thick-
nesses of adjacent dark and light laminae (La/Li), pooled for all specimens, compared with
data for Russian conophytons; (j)—Frequency distribution of the coefficient of thickening
of laminae in the crestal zone. compared with Russian conophytons; (k)—Comparison of
contour diagrams of the frequency distributions of dark and light laminae (contoured in num-
bers of readings),
93 W. V. PREISS
crystallized specimens to have proportionately
more numerous thicker Jaminae (actually thin
macralaminae), Le. the mean thickness should
be higher than for less recrystallized ones. The
following table Compares meun thickness (in
mm) af L, and Lo with degree of recrystalli-
zation observed: L. means have much ureater
spread about the total mean then L,, and the
highest means of L. correspond to the most
recrystallized specimens.
Degree of
Specimen Reerystal-
number L, méan La imean lization
S214 0.128 0.073 Well preserved
$213 0.107 0.066 Slight
S278 0.145 0.065 Slight
S277 OAL 0.084 Sticht
§32 O15 0.089 Strong
$43) D.DST 0.134 Siraug
Total inealh = D6 0,085
The data for the six specimens were com-
hined, replotted, and compared with the dis-
iibution curve of the Russian Conaphyior
gargenicum garganicun. (Fig, Thy. While the
Lo curves are very stnilary L, has a higher
mode in the South Australian form (0,08 to
1.10 mm), with asceondury peak tn the mter-
val 0.04 to 0.06 mm, which characterized the
Russian form, To some vxtent. the bimodality
ig due te errars of mevsurement arising from
the judgement ul lamina thickness sehilive to
the scale of the graduated cyepiver, and to the
Presence of thinner macrolaminse ws discussed
above,
Rate Lyth,y for Adjacent Laminac; Results
from all six specimens were pooled and plotted
in intervals of 0.25. Fhe graph compares very
closely with that of the Russian form (Pig. 11),
The dain may also be represented in the form
nf « eoutoured frequency divgram of LL.
avainst Ly. The shape and position of the
maximum are compared with those of con-
toured Russian plots; they diller only in that
the South Australian form has a displaced
seoandary peak at L, = 0.08 ta 0.10 mm, L.,
= 4K 1 0.10 mm (Fig. 1k).
Coefficient of Thickening: Randomly selected
light andl dark laminue, and macroluminae.
were measured outside the crestal zone th),
then traced into the crestal vone and remen-
sured (H), H/h was plotted at intervals of
0.5, In a total of 52 measurements, the medal
value of Hh is the interval 2.0 to 2.5 (26.9% }
while only 15.5% exceed 3.5, and none less
than 1.0 occur (Fig. Ij),
Inierspaces; Interspave = Aillings — betweeir
colunms are strongly altered, consisting of
homogencous recrystallized Uolomite Seme is
of grumous texture, composed of xenotopic
equidimensional (0.005-0.01 mm) grains,
forming patches 0.05-0.10 mm in diam., set un
a §parry matrix of grain size 0.1-0.3 mm. The
only observed remnants of primary structure
are possible smull intraclasts in one specimen.
Secondary Alteration; Fracturing of laminae
is restricted almost entirely to the crestul zones
of some specimens and marginal zones ol
others. Portions of the crestal zone are more
or less breccated and recemented in place
(Fig tid). Contertion frequently occurs
within the crestal zane, Immediately outside i,
laminae wre deflesed: this and the brecciution
ate effects prubably due to compaction during
burial, The breeeiation of macroluminac into
cleanly broken fragments several imillimetres
long suggests that the carbanate was already
lithified during the deformation (Fig. 11d). Im
places, on the column margins, Jaminae
truncate underlying laminae. Whether this is
due to peneconiemporuneous crosion or ta
sliding of the overlying Jaminue during con-
paction could not be determined, but associa-
led brecciation around the colunm margin
sugeests the latter possibility. No overfolds or
diapirie structures as in Conoplyron gargani-
cum australe Walter (1972) were observed,
supporting the idea that columns were lithifted
soon after growth.
Columns and interspaces consist entirely of
dolomite. ‘The preservation of very fine lamina-
tion suggests that dolomitizuliun was probably
penecontempoerancous, All laminae are more
or less recrystallized, the dark laminac are
coarser and more transparent that ty the Rus-
sian or Western Australian forms. Recrystal-
lization may be due to the lew grade regional
melamorphism which has wffecicd the Mt,
Lofty-Olary Are. Pale laminae between dark
mucrolaminue are preferentialhy recrystallized,
emphasizing the distinctness of the macro-
laminue, These recrystallized lansinwe consist
of sparry. hypidiviopic 19 xenotepic incqui-
uniqular dolomite, of grain size UA2-0.10 mm.
The most recrystallized specimen is a fine
marble. in which dark macroluminac, approxt-
mutely 1 mm thick, contain no preserved in-
ternal lamimac. and consist of xenotopic equi-
dimensional curhonate with interlohbate crystal
boundaries, of grain sizes 0.02-0.05 mm. The
grain. size uf the light laminae is (450.10
min, und in places much coarser, One speci-
Men is extensively sillcthed. Silicification pust-
SOUTH AUSTRALIAN STROMATOLILES I! 95
dates the prowth of the whole column, and
may be related to tectonics and diapiric em-
placement rather thun to sedimentation, Silica
consists of xehotopic quartz aggregates, of
grain size 0.05-0,10 mm, in places containing
small dolomite rthombs. Portions are com-
pletely redolomitized,
Comparisons:
Fhe conical Jamination with thickened
crestal zone, and the absence of branching dis-
ainguish this stromatolite from all groups ex-
cepl Conephyron. Tt diflers from most cono-
phytons in that the columns are not always
parallel, hue theif original growth orientation
is not clear. due to structural disturbance. On
microstructural features, it falls into the Carp-
plvios garvanicwn subgroup (Conophyion
gosanicum, C. miloradovicl, C. guuhitza, and
perhaps, ©. /aseliicuna Walter), The closely
allied C, miloradovict has more irregular and
lenueculur laminae. ©. basakicum Walter also
has very thin smooth continuous laminae, but
lacks (he distinctive Types Tt & LIL erestal zone.
The absence of numerous knotted lenses and
sharp swellings distinguishes it from the variecly
Co wirganicun nerdiciun:. On these feutures it
was assigned to Conophyieit cf. garganicus by
Glaessner er al. (1969).
The statistical study confirms the identifica-
tion as Cy garganicum garganicum, and distin-
guishes it clearly from C, gareanicion australe
Walter und C. earganicum nordicum, The
modes of thicknesses 1, and Ly. most closely
resemble C, gurganicum garganicum, especially
Ly (mode at 0,04 to 0,06 mm), while most
other conophytens have modes at mugh higher
values, C. garganicum nordicum has a modal
value of L, at O10 mm, and C. gargeretnt
ausirele Walter at about 0.08 ram. The ratio
L/L, ts the most distinctive character for
Conophyion garganicunt garganieuin. The
modal value is the interval 0.50 to 0.75, which
fulls within the broader peak of the Russian
form (0.50 to 1.00), but distinguishes it from
C. garganicum nordicum Komat, Raaben &
Semikhatov (mode 2.00) and from C. gur-
gunicum australe Walter (1.0 10 1.5), The co-
efficient of thickening is less distinciive: the
mode at 2,0 to 2.5 does not distinguish C. gar-
gonicum garganicupr, C, miiloradovic? and C.
eviindricum but excludes C, garganicui nor-
dicum and probably C. g. atesrrele.
Distribution: Lower Subsuite of the Yusmas-
takh Suite of the west and east slopes of the
Anabar Massif; the Kyutingdin, Arymas and
Webengdin Suites af the Olenek Uplift; the
Gonam Suite of the Uchur-Maya Region,
Ust’-sukharin Suite of Western Priver-
khoyan'ye, Mongoshiit Suite of the south-
eust part of the Eastern Sayan, Bul'bukhtin
Suite of the Baikalo-Patom Mountains, Sat-
kin Suite af the Southern Urals: in pre-upper
Burra Group sediments, Paratog Diapir, S.
Aust.
Age: Early and Middle Riphean: in S. Aust,
itas assumed to he early Adclaidcan-
Group GYMNOSOLEN Steinmann
Gynineselen Steinmann 1911: $8 Semi-
khatov 1962, 219. Krylov 1963: R4, Kamar
1966: 88. Krylov 1967; 346, Raaben 1969;
73 {in parti. Glacssner, Preiss & Walter
1969; LOST.
Type Fernt. Gyinitesalen eanisoyi Stein-
mann, from the Dolomitic Suite of Kanin
Peninsula; also widespread in the Southern
Urals, the Polyudey Mountuins, Kildin
Island, and Tien-Shan, USSR.
Diagnosis: Smooth to gently bumpy, swelling
amd constricting, Walled columns with frequent,
>-patallcl, often multiple branching, less fre-
quently slightly divergent branching,
Contear: Grrinesolen camsayi Steinmunn;
G. levis Krylov; G. furcunts Komur: G.
altus Semikhatovy, "“G confragesus” (in
part) Semikhatoy and G, asyminetricus
Raaben, Rauben (1969) has included part
of the group Minjaria Krylov in Gymno-
salen, chiefly on the basis of microstruc-
tutal similarity, but Krylov (1963) has
clearly distinguished Minjaria from Gym-
eesoler by its regular, subcylindrical
shape of columns, of constant diameter,
and selatively rare and simple branching.
Ape: Late Riphean,
Gymnosolen cf. ramsayi Sicinmann
FIGS, 2b-z, 3a-c, 9b, 10b, 12b,c, 134
Gyninesolen sp, Glaessnet, Preiss &
Walter, 1969: 1057.
Material: Five spegimens from nears Wilsan
Description
Mode af Occurrence: All specimens ate bould-
ers from conglomerate and breccia bedks within
the Tapley Hill Formation on the flank of a
small diapir, Only one specimen shows com-
pletely separate, discrete, vertical columns, and
is interpreted to have been derived from the
central portion of a bioherm (Fig, 13a}, OF
two specitens showing much coalescing and
bridging. ome alsa has markedly inclined
columns (Kip. 12c). These ate considered to
96 W. V. PREISS
represent the marginal portions of biohernis.
The provenance of the boulders has not been
determined.
Column Shape and Arrangement, Columns
straight lo gently. curved, erect, 1-5 em diam.,
with gentle swellings and constrictions (Fig.
2b-g). Transverse sections generally circular
to oval, but lobate and rounded-polygonal at
branches (c.g. Fig. 3a). Length of columns
hetween branches 5-20 cm. Sone columns
short (only 2-5 cm long), with rounded or
pointed terminations (Fig. 2c.e). Colunins pre-
sumed to be marginal in bioherms are inclined
(as inferred Irom the asymmetry of Jaminac
and occasional interspace Jumination). “lhe
gross morphology of marginal colunins differs
only in their frequent coalescing und bridging.
und narrow interspaces (Fig. 3b). One speci-
men with appurently erect columns is markedly
humpy (Fig, 2d).
Branching: Slightly divergent to @- or. most
commonly, y-parallel. The column expands
SA. Déeparrmenr af Mines
Fig. 2. (a)—Conophyton garganicum garganicum, from dolomite raft in Paratoa Diapir. Part of a
farge column illustrating margin structure. $528: (b) to (g)—Gyrnasalen cf, ramsayi, from
boulders in a conglomerate in the Tapley Hill Formation, near Wilson; (h), {c), (e), GF).
(g)—Vertical columns interpreted to be derived from a binherm centre. S388: (d)—Poorly
presetyed vertical columns. $390,
SOUTH AUSTRALIAN STROMATOLITES "?
rapidly, then branches suddenly into two, three
or four columns, some branches lerminaling
as pointed projections. Even in the disercte
specimens, adjacent columns muy vecusionally
coilesce. The inclined-column specimens
branch similarly, hut widening of a columo
before branching js more marked. In these
specimens adjacent hranches either are [re-
quently linked by massive bridges, or coulesce
Margin Stevervre: The surface of columns
beurs low, rounded bumps, 1 to 2 cm wide.
with a reliet of « few millimetres, Short, trans-
verse Ur inclined tihs ure exceptional, Mostly
it is covered by a wall, up to 3 mm thick, com»
pasd af from one or two to ten laminac
(Pig. 12h. cb. Generally, the marginal zone
of columns is recrystullized, but in places, la-
minue bend down near the column margin and
extend parallel to it [nr up to 2 em. Leven
where wall reerystallized. outer lamina sharp,
well preserved, In places, an tnlaminated sel-
vupe, up to 0.5 mm thick. lines the column
surface. This post-chites the wall formation,
and pre-dates the interspace sediment. In the
dixerete column specimen. bridges rare: ocesn-
sionally where two columns are closely spaced,
afew laminae may bridge ucross. Rarely, over-
hanging peaks occur: especially iF draped over
gn adjacent intractast: some columns arise
from laminae grown over intraclasts. Columns
in the inferred marginal specimens purtly un-
walled: Jaminae thin and wedge out, forming a
amuoth murgin, but do not-extend over il.
Lusaina Shape: Varies within broad limits.
Gently convex laminae most frequent. varving
in sectiod from rectangular to hemispherical,
Frequently, laminae develup two oor more
crests pror to branching, but in some cases,
incipient branches are immuuately bridped
aver. und growth of the original coJumn re-
sumed (eg. the column on the right of the
photovraph in Fig. 12b). Different lamina
shupes. occur close together in w column, ie.
degree ol inheritance of shape is low. Fig. 96
illustrates commonly occurring shapes. OF li-
minye measured, 694% have h/d Tatio between
0.2 and 6,6, the mode (26%) being the inter-
vil helween 0.2 and 0.3 (Rig. 10b), (In deter-
mining lamina shape, the poorly visible, down-
turned marginal portions of Jaminac in the
will had to be excluded). Laminae mostly
slightly wavy, with wavelength 2 or 3. mm, and
amplitude 0.2-0.5 mm.
Micrasructure: Micrositucture extensively re-
ervstallized. Where alteration minimal, alter-
uating light and dark laminue of greatly vaty-
ing thickness form w distiner streaky miero-
structure ¢Fig. |2b). Lighr Jominue are l—
0.5 mm thick Occasional thicker livht laninae
(up to | mm) may actually be peervstallized
macrolumingae, Light laminae are ¢ontinious
ucross the coluron. but thin in the wall zone
Very rarely, they are ceuticated hy micro-
unconformitivs. They are wrinkled and wavy,
corresponding to irregularities in the Wark ta-
mite. und consist of sparry, cquidimensional.
xenotopic to hypidiotopic calcite, of grain sive
UAH-O.0S fom. Trregular patches, approsi-
mately 0,05 mm diam, are stuned With a pate
brownish (organic?) pigment. Dark lena
are 0,050.3 mim thick, hut pinch und swell
rapidly along their length. In many places, they
are lenticular, consisting of contiguous lenses
or nodules 0.1-O.5 mm fang. Usually dart
laminae persist ucrass the column, but occa-
sionally Jens gut completely. so thit adjacent
light laminae merge, The clirk laminae which
ure thickest in their crests. consist of brown
pigmented xXenotopic, cauidimensional calcite,
of wrain sree O.005-U.0ES mim. In places, dark
Jaminac limonitic. In arcas of more pervasive
recrystallization, prumous textures ie deve-
loped in Which clotty remnants of dark Liminae
are set in a matrix Of sparry hypihiotopic cal-
cite. Poorly differentiated macrolaminue, 0.5-
200 mm thick, consisting of up lo & light
dark Jamination pairs, occur in many purts of
columns, The internal structure of these 1s
often not preserved, resulting in more or less
homogeneous thick dark lammae with wavy,
sharp, upper surfaces.
/nterspaces 2 mm—5S.em wide: where colunms
more widely spaced, interspaces Filled with silty
intrami¢rite. Intfachiuts are flat pebbles [5-3
em long (Figs, 12b, c; 134, suhrounded, vari
ously oriented. und foosely packed (matrix
supported), Many stand vertically in the inter
spice. Some intruclasts are curved, suggestive
of a mud-cracked origin, The matrix consists.
of broadly laminated silty, recrystallized lime
mud) fine Jaminaé, 2-5 mm thick, consist of
yenolopic calcite of grain size 0,003-0,01 mim,
while coarse faminac, of about the same thick-
ness, consist of hypictotapie 0.03-0.05 mm
frain size calcite, with much subangutar quartz,
sill, Laminations of the interspace sediment
ubut against the column walls, having accu-
mulated after the development of significant
relief of columns,
Secondary Alrerution: laminae are extensivety
altered especiully ut the marginal wall zone.
In places the lamination is completely disrupted
OB Ww. V. PREISS
ufound centres of reerystallization, but cam-
monly faint lamination or sows of dark clots
ure preserved, to indicate the presence of ori-
ginully continuous dark laminae im the wall
vone, The outer few millimetres of columns
are ocommunly recrystallized to career,
twinned hypidiotopic calcite, of grain size up
to 0.3 mm with inclusions of dark lamination
relics, In places, am acicular texture is deve-
loped in the wall zone, perpendicular to the
column murgin. The central parts of columns
ure less affected, but even here, laminuc ure
commonly reduced ta dark clots in a sparry
culetre matrix. Dolomitization of both inter-
space and columns iy found in some specimens,
where anhedral to subhedral rbombs of dolo-
mite, 0.02-0.06 mim in grain size, are scat-
tered more or less uniformly throughout a re-
crystallized sparry calcite mosaic, Frequently,
lenses of coarsely crystalline, clear calcite
occur within the lamination. Coarsely sparry
patches, cutting gcross all earlier structures, are
probably infillings of solution cavities, since
they are closely associated with discordant
stylolites. Stylolites are rather rare, and of two
génerations, The first are concordant with la-
Minae, and contain concentrations of limonite.
These are cut by major calcite veins, which in
turn are offset by the discordant stylolites men-
tioned above.
Comparisons
The stromatalites ure assigned to Gynima-
salen on the basix of their column shape,
frequent y-parallel and slightly divergent
branching, and wall, In overall column shape
an Llype of branching, presence of pointed
projections, shape of laminae and micrestruc-
ture, (he South Australian form closely re-
sembles the type G, raniwayi. Slight differences
include unwalled patches. of columns, occa-
sional peaks and hridges, and in places it
slightly bompier margin structure, G. cf, ravn-
sayi as distinguished from G_ furcatis by the
absence of markedly y-parallel, multiple
branching and by the presence of pointed pro-
jections, and from G. levis by its more widely
spaced, Icss markedly bumpy columns, G. wits
Semikhatov has apparently been affected hy uw
strong cleavage, and its columns are slightly
deformed, making comparisons difficult, but it
appears to have a more continuous, banded
lamination. G. aywinmetricus Raaben has thin-
ner, smoother laminae thin G. ramsayi. G.
confragoxus Semikhutov has in part (specimens
from the Shorikhin Suite) deen reassigned by
Raaben (1969) to Inzeria Ul. confraeosa);
ese specimens are distinguished from G
ranisayl by their irregular columns, interrupted
will and more frequent peaky and cornices.
Semikhatoy's specimens from the Dashkin
Suste, now considered as Vendian (Krylov i
Rozanoy er af. 1969, p. 215), have much
smaller, bumpier columns than G. retmuseeyi.
Distribution; Sub-Inzer Beds of the Katay
Suite und Minjar Suite of the Karatau Series
of the Southern Urals: Niz’ven Suite of the
Polyudov Mountains: Carbonate Beds of the
Metamorphic Series of the Kunin Peninsula:
Kil'din Series of Kil'din Island; possihly the
Spatagmites af Norway; Bystrin Suire of
Southern Timan; Chatkaragai Suite af ‘Ten-
Shan: as clasts in Tapley Hill Formation. &
km E of Wilson, southern Flinders Runes.
S$, Aust.
Age: Late Riphean, in S. Aust,, not younger
than the Tapley Hill Formation.
Group INZERIA Krylov
Mngeria Kryloy 1963: 71. Krylov 1967;
29. Cloud & Semikhatov 1969; 1042.
Ravben 1969: 77, Glaessner, Preiss &
Walter 1969: 1057.
Type Form: Inzerid tianusi Krylov, from
the Katuv Suite of the southern Urals,
and the Demin Suite of the Polyuday
Mountains, USSR,
Diagnosis: Subparalicl, usually wnwalled, sub-
cylindncal, ribbed columns, frequently with
niches containimg projections Branching
mostly a- to @-parallel to slightly divergent.
rarely y-parallel or markedly divergent.
Conn: Lf. gomasi Krylov, £, toctagnlli
Krylov, f. intia Walter, and probably /,
djejimi Riaben and fF, ayfryslandica
Raaben, 4 (Minjarvia) nimbifera Semi-
khatov may he included, but Raaben
{1969u) has placed at in synonymy with
i. fjomusi, and has partly reassigned
Gynmosolen canfrayosus —Semikhatov
ta Snzeria (L eonfragoxa), Raaben haus,
however, considerahly broadencd the con-
cept of J/rseria, placing lille importance
on Krylev’s (1963) critema of ribbed
columns with niche-projectiony, On this
basis, Aldania Kryloy (ia Rozanov e¢¢ ol,
1969) could perhaps be better included in
fnzeria. Descriptions of 2. macula, 0.
varinsata, J, sevintedr and J. chunitbergiva
Golovanov were unavailable, but Raahen’s
(1969, Fig. 2L) illustration of frzerie
nrteuld docs not resemble any other des-
eribed J/nzeria. The new S. Alist, forms
are I, multiplex und |, conjpimera,
SOUTH AUSTRALIAN STROMATOLITES iL ed
Age and Distribution: \ate Riphean, wide-
spread in the USSR: Bellon Springs. Forma-
tion. Central Aust.; Brighton Limestone and
Wundowie Limestone, S, Aust; Hinde Dolo-
mite. and doubtfully, Dook Creek Forma-
lun, NT,
Inzeria cf. tjomusi Krylov
FIGS, 3h. 2, tieg, |3b-
Material: Three specimens from Burr Well.
Description
Made of Occurrence: “Vhe stromatolites form
a lenticular bed, interbedded in green shales,
and consist of fottr or five contiguous gently
damed bjioherms (Fig, (3c), 2-+ m diam,
with a inuximum thickness of about $ mi. Toe
wirds the west, the bed thins and lenses nut
grsdually; in the easterly extension, columiar
stromatolites give way to flat-laminated lime-
stone. The lower partion of a domed bioherm
consists of Hat-laminuted stromatolitic lime-
slone. or contiguous, very broad cummuly (part
of which are seen in Fig. 13d). up to 20 em
thick; overlying this (hut never seen in sedi-
mentary contuct with it), is a zonc, up to 20
em thick, of discrete, verticul, subcylindrical
eolurons, 2-10 em Wide, The base of these
columms is an intensely stylolitic zone, in which
aw thickness of up to several centimetres has
been removed by solution (Figs. 13b, d). At
the maruins uf bioherms, the columns become
inregular and slightly inclined from’ vertical.
Columns jre bridped over hy a thin, poorly
exposed zone of flat-laminated stromatolites.
Column Stupe and Arrangenenrs; Columns
short, subeylindrical, with some swellings and
constrictions, of diameter 2-10 cm (Figs, 3f. 2.
4a-g), henght 10-20 cnt (the whole thickness
of the columnar zone}, Transverse sections
round, rounded polygonal or slightly lobate.
Columns have vertical, straight axes in the
central parts. of bioherms, hut hecome irregular
at the edges.
Branching into discrete new columns rare, per-
haps due to the small thickness of the bed.
Niche-projections very frequent; they are short,
narrow, usually rounded, sonelimes xlighthy
elongated, set into niches in the side of the
tain column, which, most commonly. resumes
its former diameter at the top of the niche
4Figs. 13b, e: du, d, & g). Occasionally auja-
cent columns coalesce,
Margin Structure; Due to strong recrystiliza-
ton of columns the margin structure is ob-
secure. Laminue upproach the margin al u high
angle. and are not defiecxed at their cdges;
columns always unwalled. Lateral surface of
columns with numerous short transverse vibs,
up to 2 cm long, occysinnyl overhanging la-
ininue and peaks. Th places, adjucent columns
linked by bridges up to 5 mm thick,
Luminae Shape: Always genlly conver, vary-
ing in shape from continueusly curved domes
la very low, obtuse cones. as illustewted in Fig.
9c. Lamina shape inherited from underlying
Jaminae. without fapid chynves in convexity.
Ratios of h/d usvally low; 91% of laminae
huve h/d between 0.7 to 0.4 (Pig. 10e). Fine-
seule structure of laminae smooth to gently
wrinkled.
Microstructure: Strongly recrystallized through-
out columns, but in places the gross indistinctly
banded structure of laminae ts moderately well
preserved (Fig, 13e). Relatively thicker light
Jaminaé alternate with thinner datk laminae
hut recrystallization has in places obliterated
the distinction. All laminue have diffuse baun-
daties, Light laminae 0.2-2.5 mm thick, com-
monty significantly thicker al their crests than
their edges. especially in the obtusely-conical
laminae. ‘They consist of @ sparry, equigtanu-
Jar, hypidiotopic mosaic of calcite, of grain
size OES—O.02 mm, with ingluded small, irre
gular patches of darker pigmentation. Derk
laminae githee smooth or finely wrinkted
(largely due to embayment by recrystallized
adjacent light laminae), thickness 0,2-1.0 mm,
generally thinner than adjacent laminue. Oeca-
sional thinner dark laminae are lenticular, but
whether this feature is primury or due to Te-
erystallization is unresolved, Like the light In-
minae, they are slightly thickened in eheir
crests. Dark laminae consist of xenotopic,
slightly inequigranular calcite, grain size 0.005-
0.09 mm, stained with a pale brownish pig-
ment.
Interspaces filled with homogeneous recrystal-
lized lime mud, with occasional intraclasts,
Culcite xenotopic to hypidiotopic, grain size
0,01-0.03 mm, contains about 5% angular
quartz silt, Quariz grains corroded by recrys-
tallized calcite. Occasional flat intraclasts up to
1 cm long, with diffuse boundaries, in parts
of interspaces, recrystallized to sparry calcite
mosaic, prain size 0.02-0.03 mm-
Secondary Alteration: The whole reck is per-
vasively altered, While columns are pale grey,
tunsparent in thin section (Fig, 3c). the dark
laminae perhaps tinted with arganic milter,
iterspaces are pale buff probably due to the
presence of small amounts of limonile, Neither
columns nor interspaces are dolomitized, The
100 W. V. PREESS
S.A DérartmaAns of mines
FIG. 3
SOUTH AUSTRALIAN STROMATOLITES UI IU}
boundary between interspace and column is
always diffuse, obliterated by recrystullization
in both, This reduces the reliability of the re-
constructions. Highly irregular styloliles with
large Jubes sepatale the basal laminated sedi-
ment from the discrete columns, with a gone
up to 5 cm thick of intense brecciation and
late-sisge infilling of fractures by coarsely
ervstalline calcite. Possible remnants of the
lower portions of columns, highly enriched in
limonite, are sometimes preserved between
cross-cutting stylolites (Fig. 13e), Large sub-
aphericul nodules, up to 3 cm diam., of
coarsely crystalline culcite are very commun
in the litnestone at this locality. mostly located
withm columns, Twinned calcite crystals in
these highly elongated, 1-3 mm wide, up to
3 om long, vertical or radially arranged. Mast
crystals terminated upwards; their acute tere
minations project into the laminated limestone
of columns, The major cross-cutting stylolites
post-date the coarsely crystalline nodules. Qver
large areas, columns are completely tecrystal-
lized so that lamination is partly or totally ebli-
terated, Such areas consist of xenotepic, to
hypidiolopie mosaic calcite, gtain size up to
OS mm. Where rtecrystallization incomplete,
regular fragments of disvapted dark laminae
surrounded by sparry, recrystallized mosaic
calcite.
Caniparisons
The presence of ribbed columns with nu-
merous niche-projections places the stromato-
-lites in the group Inzeria, They are differen-
tiated from all other Australian forms of
fazerla and from /. rocrogulii Krylov and &
djejimi Raaben by their very infrequent
branching, consistently gently convex laminae
(grading to low-conical rather than reclangu-
far}, and their short Jength of columns. In hav-
ing subeylinerical, erect, mbhed columns with
nunverous niche-projections, they closely te-
semble Russian specimens of /, tjorud Krvlov,
hut differ in the thinness of the zone of
columns; the dbsence of branching may simply
be a consequence of the short length of
columns, Unlike 7, tjeaiuest [rom the Southern
Urals, steeply convex laminae arc absent. The
microstructure with pinching and swelling or
wrinkled dark laminae is similar, but the pro-
minent concentrations of iron oxides along
Fig,
concordant solution surfaces are ahsent. Until
bicherms are found in which the columns had
the opportunity to grow to 4 greater height,
so that the mode and frequency of branching
can be determined, and which are less recrys-
tallized. so as to preserve the margin structure.
no reliable identification ts possible,
Distribution: Middle limestone band of the
Wundowie Limestone Member, Umberatuna
Group: Burr Well, northern Flinders
Ranges, 8. Aust.
Age: Late Adelaidean, correlated with the
Late Riphean or Wendian of the USSR,
fnzeria conjuncta £. nov.
FIGS. 4h-m, 9d, 10d. 144, b
Muterial: Three specimens from near Depot
Creek.
Holotype: S402 (Figs. 4a, b, tj, 14a), from
the Brighton Limestone equivalent, 3 km
north of Depot Flat H.S., southern Flinders
Ranges, 8, Aust,
Neme: Latin cenjurea, meaning joined,
refers to the frequent coalescing and bridg-
ing of columns,
Diagnosis: Inzeria with broad, unwalled, rarely
branching, frequently bridged und coalescing
basal columns, which divide by «-parallel
branching into narrower, unwalled upper
columns with occasional e- and -parallel
brinches, Niche-projections moderutely fre-
quent, Laminae nearly flat to rectangular or
gently convex, wavy or wrinkled, with a dis-
tinct streaky microstructure.
Deseriplion
Made of Oeveurrence: Fiekd examination of
bioherms is hampered by the very extensive
lichen cover on rock, faces and by the discon-
tinuous outcrop. Three domed bioherms, up
to 50 m long, 3 m thick, occur interbedded in
massive oohtic and intraclastic limestone, and
cannot readily be differentiated from Acaciella
augusta in the field, The basal, central portion
of bioherms consists of flat-laminated stroma-
tolitic limestone, passing up gradationally into
bread columns, 5-20 em wide with frequent
coulescing und massive bridges. Flat-laminated
intervals may. intervene, At slightly different
levels, the broad columns divide by a-parallel
branching into 1-5 em wide columns.
3. (a) to fe)--—Gymiasolen cf. ramyay: from near Wilson, (&#)—S388, vertical columns pro-
bably fron: a bidherm centre; (b) and (d)—S387, inclined columns interpreted to be derived
from iw bioherm margin; (¢)—S389; (¢)—S390, irregular and frequently coalescing columns,
Traced from slabs; (f} and (g)—Jnereria ef. tienes’. Wundowie Limestone Member, Burr
Well, northern Flinders Ranges. $479.
102 W. ¥. PREISS
S.A, Department pf Miaies
Fig. 4. (a) ta (g)—lnzeria cf. umuasi, Wundowie Limestone Member, Burr Well, Northern Flinders
Ranges. (a). (b) and (cy, $5942; (d). (f£), and (2)—S480: (c)—S479: (fh) to Gin)—lazeria
conjuncte, Brighton Limestone equivalent, Depot Creek; (h), (i), and (j)—Holotype $402,
broad basal columns branching into narrow colunms; (k)—S403; elangate columns from a
bioherm margin; (1) and (m)—S398; narrow upper columns.
SOUTH AUSTRALIAN STROMATOLITES I
Caluma Shape und Arraigeneiu: Broad
columns in lower part of biuherms subeytin=
drical, up to 30 em high In their discrete por-
lions, comMronly with rounded polygunal trans-
verse sections, Where adjacent columns
coulesce, or a wider columm branches, transvetse
sections may be complesly lobate. Gverlying
narrow columns slightly clongated, from 1 x 2
cm ty 3 x 5 cm in transverse section, up to
15 cm long between branches. Columns within
central part of bioherm straight. erect (Fig. 4h,
i,j) while at margins, they become inclined at
45°. slightly curved and strongly elongated
(Figs. 4k, 14b), with swellings and constrice
\ions
franchiitg: Niche-projections are formed by
unequal, o-parallel branching, or, less com-
monly, divergent branching; the narrower
column is set into the niche in the main wide
column, Which generally resumes its former
diumetec after the termination of the projec-
lion, Where projections branch divergently,
they protrude beyond the margin of the main
column. Projections U.5=4 em tong. Within
hroad column level, branching (other than by
niche-projections) tare, Broad columns then
divide by a-parallel, rarely §-parallel branch-
ing, into narrower 1-5 cm wide columns,
which hranch again, less frequently, by o- or
#-parallel branching, In marginal zone of hio-
herm, branching #- to +-pafallel, often with
constriction at branching; niches stil] common,
but clongated parallel to the long axes of platy
colimns (Fig. 4k),
Margin Structure: Lateral surface uneven, with
very frequent. transverse ribs. same small pro-
jection’, bumps, bridges, und accasional small
peaks, Ribs, 0,5-1 cm wide. may be followed
around the column periphery for a few cenu-
metres. Both massive and delicate bridges accur
between adjacent colunins, and, sametimes,
between columns and projections, Niches in
the column margins 4 to several centinrebres
deep: some niches partly cloyed at one end
(Fig. 4h). Occasional niches clongated trans-
versely, grading into prominent ribs (Fig. 4h,
ji. There is no wall; at the column murgins,
laminae thin oanty slightly, and either end
abruptly or turn down and wedge out; they
do not envelop the lateral surface of the
column f{Fig. t4a).
Lamina sfape varices greatly from broad
columns to upper narrow columns. In broad
columns most laminae flat, geally convex, or
reclangular (Fig. 9d). In plices laminae deve-
lop to of more ¢rests, then either the column
103
branches (if near the branching level) or the
Interspuce is bridged over. and the column re-
sumes its normal growth (Fig. 14a}. In broad
columns, measured values of h/d ore below
O25. In the nyrrow, upper columns, laminae
ure Consistently more steeply convex. OF those
measured, 81% lic between 0,3 and 0.4, Col-
umns in the marginal zone of hioherms have
laminae strongly asymmetrical towards the ex-
terior of the bioherms, commonly as steeply
convex as in the upper narrow colamns ( 60%,
of h/d between 0.3 wml 0.4). Fig. Md illues-
trates the distribution of lamina convexites. All
laminac wavy, with wavelength 2—+ mm, and
locally wrinkled.
Microstructure distinctly streaky with butt
lenticular and continuous, wavy, swelling and
constricting laminae, Dark laminee O,05-0,3
mm thick, wrinkled and wavy, with non-paral-
lel upper and lower boundanes, sometimes
atading into aligned clots and lenses, They
consist of equigranular hypidielopie 10 idio-
topic dolomite, grain size 6.005-0.0L mm.
Crystals equidimensional and stained a pale
green lint, responsible for thé green colour of
the luminue. No individual grains of pigment
could be resolved even at 1200 x magnifica-
tion. Amplitude of waves and wrinkles 0.2-0.5
rom, wnd thickness of Iaminae changes rapidly
within a few millimetres. Ligh leniinae consist
of white to pale grey parily dolomitized sparry
calcite of axenotopic to hypidiotopic texture,
grain size 0.005—0,035 mm, tending to lens out
near column murging where adjacent dark
laminae merge. They also contain some coarser
detritus, including fing sund-sized, well rounded
quartz and feldspar, and small dolomite
thombs similar to those of the durk Jaminac
but less pigmented. Over most of the area of
thin sections, dark laminae tend to be grouped
into: macrolaminae 1-5 mm thick, which, like
individual laminae, pinch and swell murkedty-
There is evidence of minor contemporaneous
erosion of thickenings and waye-crests of
macrolaminze.
Interspaces; Both lower broad and upper uar-
row columns are separated by narrow inter-
spaces, 1-20 mm wide, but columns from bio-
herm mur'gins are almost in contact. The infill-
ing sediment ts layered, either by sandy lu-
Minac, or by single stromatolitic laminae
bridging between columns, Interspace laminae
ate depressed, concave upwards (Fig. 14n?.
‘The carbonate of interspsces is a dolemitized
limestone: slightly inequigranular hypidiotopic
calcite (partly recrystallized lime mud), grain
PREISS
WwW. Y.
104
wy
SA, Depareoyern at tu
FIG. 5
SOUTH AUSTRALIAN STROMATOLITFS I
size 0.005-0.02 mm, contains subhedrul
rhombs of dolamite, 0.005-0.05 mm diam, In
places, quartz sand laminoe up to a few mil-
limetres Thick abut against the column margins,
suggesting that they postdate the growth of
that portion of the adjacent column. No cur-
bonale allochems were observed Av times of
bridging, the structures had a relief of less than
une centimetre, und bridging laminae may be
only one or two centimetres apart.
Secondary Alterqrion: Quartz and feldspar
erains, oth in columns and interspace sedi-
ment, have corroded bounduries; in pluces their
margins ate completely replaced by carban-
ate. While the dark laminae ure almost coms
pletely dolomitized, the lime mud comprising
the light laminae and the interspace filling is
only patchily dolomitized and also contains
hypidiotopic, coarser culcite due 1 partial re-
crystallization. The dolomitization is probably
secondary. Stylolites rare cacept in the bioherm
margins, where they separate contiguous
columns, Smull vughs. up to 3 mm diam, filled
with course, twinned sparry calcite occasiooully
cut across the lamination. The origin of the
green coloration of dark Jaminae is not clear,
since no particles of pigment could be resolved:
the dolomite crystals themselves are tinted.
Sunuce oxidation during weathering cither
partly removes the colour, or depasils yellow-
brown. limanite in interspaces or along style-
lites.
Comparisons
The stvomatolites are difficult to distinguish
in the field tram Acuciella cuynsta Preiss, but
ure assigned le Jazeria on the following charac-
ters! ribbed lateral surface, absence of wall,
dominance of purallel branching, and niche-
projections, The upper narrow columns also
resemble Ketaviie and Kulparie but are dis-
tinguished by the presence of long transverse
ribs, the absence of a wall. and by microstruc-
ture; unlike Katavia and Kulparia, their pro-
jeettons are usually rounded, und set in niches.
7. conjuncta differs fram 7, tlamusi Krylov and
/. mie Walter in having frequently coalescing
columns, and consistently genily conver, wavy
and wrinkled laminue; it lacks the consistently
elongated niche-projections and the complex
bioherms of /. intia. Unlike I. djejiri Raaben,
its columins are straight, with frequent niche-
Ins
projections, and rarer branching. J. conjuttcte
is distinguished from /. toctogulii Krylov by
ily less frequent, dominantly o-parallet branch,
ing, and by its coalescing and bridging, J. con-
juncta is especially similar to Aldania sibirica
in margin structure and mictustructure, but
has more irregular and coalescing columbss. As
pointed out above, A/dania night he hetter
included in Jazeria.
Distribution: Brighton Limestone equivalent,
3 km north of Depot Flat H.S.. southern
Flinders Ranges, S. Aust
Age: Late Adelaidean, correlfted with the
Yate Riphean of the USSR.
Inzeria multiplex f. nov,
FIGS. 5, Ge, 1Ne, 14c, d, Sy
Material: Six specimens froai figal Melrose
and Yednulue.
Holotype; S385, from the Brighton Lime-
stone equivalent, 8 km NW of Mt, Remark-
able, near Melrose, S, Aust. (Figs, Sa-i, 14,
d).
Name; Latin rudtiplex, meoning complex,
manifold or with many parts,
Diagnosly: Inzgeria with Trequent, dichotomous
to multiple, a- and A-parallel to slightly diver-
gent branching, and rarer branches arising
from niches, Columns have irregular transverse
sections. Margin bears ribs. bumps und shor
projections. Laminae gently convex, smanth to
wrinkled, with regularly streaky microstvuc-
ture.
Deseriprion
Made of Occurrence: Due to poor outcrop,
the exact mode of occurrence at Mt. Remark-
able is not known; a kirge bioherm is inferred,
since, when followed along strike, the stroma-
tolitic bed passes into massive intraclastic
limestone, but the contact is not exposed, Al
Yednalue, the stromatolites form a very thick
bed, which has not been waced Juterally. In
outcrop, the stromatolites at Mt, Remarkable
resemble literally linked forms, and columns
becoine discernible only when the rock surface
1s Cut.
Column Shape and Arrangement, Colunnis
tuberous to subeylindrical, erect to inclined
(Figs. 5; 14c, d), with steaight or gently curved
axes: Occasional columns sharply bent, espec-
tally when ussoci:ted with coalescing, Height
Fig,
*, (a) to ())—fazeria multiple, Brighton Limestone equivalent, southern Minders Ranges:
(a-i)—Holotype $385, west of Mount Remarkable: ¢j, k}—S499. Flogt specimen, east of
Yednalue, (1, m)—S498_ Outcrop specimens, cast of Yednalue.
hits Ww.
of columns between branching 4-21) emt, Trans-
verse sections of columns round or rounded
polygonal to irregular and lobate at points of
branehing or coulescing. Columns which may
be variously clongated, vary From 1 lo 5 em
in diam. AL top of bed, columns frequently
bridged by continuous, laterally linked layers.
Branching very frequent and complex, either
arising (fom niches in the parent column
{Fig. Si). or, most commonly, by equal divi-
sion (Mig. 5a, b, ¢), usually 2-parallel. rarely
u- OF y-parallel, or slightly divergent (Figs. 14c,
H5a). Adjacent columns frequently coalesce,
especially in the upper part of bed,
Marein Srructare; Column margin’ irregular,
with numerous, short transverse ribs, low
bumps and some Slizhily overhanging laminac.
Bumps and ribs tocally grade into yery short,
outgrowing projections, less than 1 cm long,
which are more common than projections set.
in niches, especially in the Mt. Remarkable
specimens (Pig, 14d). There is no wall; com-
monly gently convex laminae terminate at the
column margin, without bending over, some-
times overhanging to form sniall peaks and
cornices. Small pertions of column margins
relatively smooth. Bridges involving any num-
ber of laminae are common, especiully near top
of bec.
Lanning Shape: Almost ulways gently convex
(Fig, 9e}) even in the narrowest columns h/d
docs nut exeeed 0,5, Ol laminae measured,
93% have h/d between 0.1 and 0.4, the made
(40%) being in the range between U2 and
0.4 (Rig. Tey. Larminac may be dayhbly-
crested, prior to branching, On a small-scale,
laminag brondly wavy, and in places slightly
wrinkled,
Microsiructure: Alternalion of light, sparry
laminae and dark, iron-stained laminae, with
indistinct boundaries and varying continuity.
In places, laminae grouped into macrolaminae
1 or 2mm thick. Boundaries between laminac
frequently wrinkled. Light laminae (0.1-1.5
mm thick, usuilly constant weross the column
width, smooth, wrinkled or wavy (Fig, 14c),
with parallel wpper ynd lower boundaries.
Varying abundances of fine: quartz sand and
silt are incorporated in the light laminae, which
consist of hypidiotopic to idiotopic carbonale.
erain size 0.01-0,03 mm. Grains equidimen-
sional, sometimes cuhedral. Dark laminae
thinner, generally 0,1-0.5 mm, but pinch and
swell geross the column width; crests wl
laminae commonly thickesi Dark taminac
V. PREISS
erade [ron smooth fo wrinkled, and frequently
become discontinuous, forming chains of clote
and lenses up to 1 mm long, separated hy
sparry carbonate (Fig. I4c), Dark laminac.,
clots and Jensex composed of reddish-brown
iron-stained, xXenotopic carbonate, grain size
0.003--1.01 am,
liiterspaces: Columns gencrally closely spaced,
interspace width 1 mm-—* cm, The sediment es
different in the two areas of occurrence.
t1)At Mt. Remarkable, if is broadly lamnu-
leU redilish dolomicrile; laminuc 1— mm thick.
generally fist or slightly concave upwards
Darker laminae generally thinner (up te |
mm), of xenotepic dolontite of yrain size
O.N03-—0,005 mm, alternating with thicker,
paler, laminue, up to 4 mm thick, of xenotepic
dolomile, 0,005-0.015 mm grain size, with a
high pereentuge of terrigenous detritus. [angu-
lar ynartz silt of grain size 0,02-0.05 mot,
und occusionu) micu flakes). Intraclasls up to
T cm Jong, 2 mm thick, locally present in
interspace. generally siunding vertically or in-
clined (Figs. l4e, d).
(2) At Yednalue, the interspaces are filled with
unlaminated sandy limestone, with quarts snd
feldspar grains, 0.1-1.0 mm grain size, sub-
rounded to well rounded, all embayed hv
hypidiotopic to idiotopic calcite cement of
grain size up to 0.6 mm, Sand grains mostly
tightly packed, in places separated by a green-
ish argillaceous matrix (Big. 15a).
Secondary Alteration, Specimens from Mt.
Remarkable consist entirely of dolomite, while
those from Yednalue are calcite. Mt. Remark-
able specimens are, however, better preserved:
the idiotopic and hypidiotopic dolomite pro-
bably formed during early diagencsis, but did
not destroy the fine structure of the stroma-
lolites. The dolomitic rock may have proved
Mare resistant to later recrystallizalion, which
has in both areas disrupted the fine damination
to a greater or lesser extent, In adthtion,
cleavage is well developed at Yednalue, and
the columns are slightly deformed, sa thit
metamorphism may partly account for the
greater reerystallization here, Occasianal con-
cordant slightly sutured stylolites follow the
lamination, sometimes affecting xeveral adja-
cent colunins, but all are cross-cutting ono fine
scale. Greenish orgillaceous matenal is con-
centrated in the stylolites. Some stylolites fol-
low column margins and thus remove the
Minor surface features of columns (Fig. 15a).
Tectonic veins are filled with quartz or calcite.
SOUTH AUSTRALIAN
Compur/sans
The stfomatelites arc assigned to Ingerta
because of their ribbed columns with projes-
lions, but they frequently resemble Baicalia in
their tuberous shape; Baicd/ia, however. much
more ollen hus divergent branching, more over-
hanging lominae, and a distinctly banded
microstructure. In having some a-parallel
branching,, they resemble Kassiella Krylov
and Acacielle Walter, but are distinguished by
their Frequent f@-parullel branching and
branching from niches, Inzeria sueltiplex 1s
distinguished from f. rjomusi Krylov, 1 inva
Waller, and /. confined Preiss by its very fre-
quent branching. and rare projections set in
niches. In these features it resembles J, focfe-
eulti Krylov and J. djejind Raaben, but /. rocte-
euiit has more regular, cylindrical colunins,
while F. djenimi has steeply convex laminae.
Distribution: Brighton Limestane equivalent;
8 km NW of Mt. Remarkable and 12 km E
of Yednaluc, southern Flinders Ranges, S,
Aust.
Age; Late Adelaidean, corretited with the
Late Riphean of the USSR.
Group JLUIRUSANTA. Krylov
Jiirusenia Kervlev 1963; S81. Raaben
L964; 93, Krylov ir Rozanoy er af, 1969:
195. Cloud & Semikhatoy 1969: 1045-
Semikhatoy, Komar & Setebryakov 1970;
166. Bertrand-Sarfati 1972: 52.
Type Form: Jurisania cylindrica Krylov,
from the Katay Suite of the Southern
Urals.
Digenoasis; Even, — parallel, — subeylindrical
columns with round ot oval transverse sections
and tare, dichotumuus g-parallel branching.
Columns partly walled, partly bear downward
directed peaks and overhanging laminac, fre-
quently covered with an unlaminated selvige,
Conen Jurusania cylindrica Krylov, J,
tumuldurice Krylov, J. nisvensis Raaben
and J, judemicn Komar & Semikhatov. J,
shirica Jakoviey has been transferred by
Krylov (1969) to a new group, Aldana,
but Semikhatov, Komar & Serebryukov
(1970) retain ils assignment to Junesanni,
Bertrand-Sarfati (1972) has erceted new
forms J. derbalensis, J, lisse, J. alte,
Age: Late Riphean to Vendian,
Jurusania burrensis f. nov,
FIGS. Ga-l, YF 10f, 15b-, 16a
Miulertalt Four specimens trom Burr Well.
Hoefotype; S543 trom the upper limestone
SUROMATOLITES 1 17
band of the Wundowte Limestone Menrber.
Wirr Well, northern Flinders Ranges, 8.
Aust. (Figs. bd. ¢, f, 15e).
Name: After the Burr River, on the bank of
which the stromatolites occur.
Diagnosis; Jurusania with smooth to gently
bumpy, partly walled columns and local, short
peaks and overhanging laminae. Lamina shape
gently convex to subconical, laminae lenticular
with diffuse, streaky microstructure. Columns
puttly covered by an unlaminuted selvage.
Descripion
Mode of Oecurrence; The stromatolites occur
in lenticular beds af contiguous spherical and
subspherical bioherms up to 2 m diam. (Fig.
L5¢c). The bioherms consist of 3 concentrically
arranged zones. capped hy an undulating
columnar zone. Bioherm cores up to 50 cm
thick consist of irregularly pseudocolumoaur
and columnar-layered stromatolites of dark
grey limestone, overlying sandy limestone with
large, reworked intriclasts. Cores surrounded
by concentrically laminated zone, from whith
long straight, parallel columns arise. At bio-
herm margins, columns slightly inclined, rarely
subhorizontal: generally columns remain sub-
parallel throughout the bioherm, but show
more bridging and coalescing al. margins,
Spherical bicherms, mutually in contact. over-
lain by Mat or broadly undulating 1 m thick
beds of columns with numerous bridges and
pseudocolumns.
Column Shape and Arrangement. Coluinns
long. straight, purallel or radially arranged,
cylindrical or subeylindrical. In one specimen
from near the base of a bioherm, columns
somewhat inclined. irregular, tuberous, und of
strongly eblipuieul op Jobate transverse section
(Fig. Gb, c); otherwise transverse sections
round or slightly ellipticul. Columns mainly
smooth, with only occasional low, hroud bumps
(Figs. Ga-d, 15h, d, e); single columns gene-
rally have constant diameter, 5-10 em for basal
columns (Fig, 6h) und 2 em for upper, nar-
row columos (Fig. 6d, ¢, £), Length of columns
between branches nwy exceed 30 em; the
whole columnar zone of bioherms is up ta 1
mi thick, Columns in the overlying undulating:
bed rather short, and {frequently bridged, ap-
parently arising trom basal, flat-laminated
siromalolites (Figs. 6a, 6a, specimen S48.
but the exact location of this specimen in the
hioherin is not certain).
Branching cather infrequent especially in nar-
Tow, Upperthost columns, which may be up lo
108 W. V. PREISS
SA. Department st Mines
FIG. 6
SOUVH AUSTRALIAN STROMATOLILES 11
30 cm long between branches, but more often
terminate their growth befere branching,
Branching wlways dichotomous, either q-. or
slightly B-parallel (Figs. 6g, b), Occasionully
two neighbouring colunms miy cualesce, espe-
cially in upper parts of bioherms.
Margin Sirucinre geverally smooth, walled or
unwalled, bearing only hroad Jow bumps
several cm wide and of telicl up to 5 mm
(Figs. 6a-h), Lamjnic generally approach the
margin ul an acule aligle, but the actual margin
is Lrequently removed by stylolites. In areas not
aftecuod by stylolites. laminae either terminate
at mareii, or extend down for u distance of up
to |ocm to form a patchy wall (Fig. 15d, e).
{ns a few places, laminae overhang slightly to
form) small peaks, a lew mn long (Pig. Ge, f,
2), Larye overhanging peaks developed only
an the irregular columns (rig. Ob). from the
lower parts of bioherms. Considerable areas
ul smooth columns coated with a xclyage, 0.2-
1.0 mm thick. of unlaminuted very fine grated
culeite 1 Fig. 15d).
Lamina Shape varies to some catent with
column width, but most laminae gently convex
with relief about | cm (85% have h/d between
.2 and 0.5, Big, LOf): # Few narrow columns
haye <teeply convex to subconical Jaminac
(Fie. 9f). Laminae smoothly curved, micro-
unconformities rare, Laminy shape always in-
herited from the underlying laminae, so that
ho marked, tapid changes occur. Fine seule
structure of lamibue lenticular and very gently
wavy, with wavelength of 2-5 min, amplitude
Oo. mm
Microsimeivre: The dark limestone comprising
the pscudocolumnir bioherm cores is almost
entirely recrystallized, and even the lamination
is yarely preserved. Lamination in columnar
parts of hioherms diffuse, streaky, consisting of
altemating, finely wavy, lenticular, dolomitized
sparry pale laminae and darker, micritic
laminac, which intergrade. Micritic lanvnae te-
crystallized to microspar, grain size 0.005—
0.015 mm, of xenotopie, polygonal, equi-
granular texture: they have very vague boun-
daries, and vary in thickness from 0.2-0.5 mm
over short distances. Tn places, luminae thin
and terminate Jaterally, or consist of short,
Fig.
hut
ligned lenses. a few millimetres long. Sperry
laminae O.1-0.5 mm thick, pinching and swell-
ing but more continuous ucross colunin width,
consist of hypidiotopic to xenolopic calcite.
crain size 0.01-0.03 mm, with scattered sub-
hedral dolomite crystals, grain size 0.015-
0.06 mm. In places, laminae almost completcly
dolomitized, consisting of closely packed hypa-
diotopic dolomite with remnant interstitial
sparry calcile, Sparry laminae may nlso be
completely recrystallized, with little dolomitiza-
tion, to a hypidiotopic mosaic of grain size up
to 0.2 mm. The unlaminated selvage present
in places on cohinin margins consists of xeno-
topic calcite mosaic, grain size 0,005-0.02 mm;
its origin is not. clear (see secondary alteration),
fuerypuces: Columns 0,5-2.0 cm apart. Mter-
spaces filled with poorly bedded intramicrite.
partiwly dolomitized. Thtractasts smostly fiat
limestone pebbles, 0.5-3 mm thick, 2-30 mm
long, generally lying parallel to bedding, or
standing vertically in narrowest inierspaces
(Fig, 15d), The flat pebbles, which commonly
have rounded margins, consist of xenotopic,
eqyuigranulur mosaic eulcite of grain size up in
0.01 mm, and contain scattered subhedril dolo-
mite erystals, grain size O.0TO-0,.0L5 mim. Sub-
taunded to well rounded quartz ond Eeldspir
sand grains occur in places. Intraclasts mode-
rately loosely packed, so that some in contact.
some not; sediment was probably matrix-sup-
ported. The matrix. probably originally micti-
tic calcite, recrystallized 1o xenotopic inequi-
granular texture, grain size up to 0.015 mm,
oceysionally with scattered dolontite erystals.
The matrix, may he preferentially dolomitized,
Tn the specimen apparently fram the unitula-
ting bed capping hioherms, interspaces filled
with markedly upward concave laminated, re-
crystallized lime mud, without Intractasts; In-
minue somewhat thicker (approximately 3
mm) and more regular than those of the stro-
matolite columns.
Secondary Alteration: Kven thé finest calcite
laminae have probably undergone some recrys-
tallization to form 4 very Tine grained calcite
mosaic, Dolomitization apparently postdates
this, as subhedral dolomite crystals cut across
the calcile mosaic. In places, especially meal
fi. (a) to ¢h)—JSurusania brerensis, Wuodowie Limestone Meniber, Bure Well, nistthern Fit:
ders Ranges! (a)—S481; basal columns arising from wndulating stramatolites; (bj, foy—
S483: irregular columns at biohermy margin; (a), (2), (£)—Holotype $343; regular narrow
upper columns; (2), Ch
Brighton Limestone equivalent,
(h)—S482; regular broad, Jower eolurans: (i) to (o}—-Kytavia costata,
Depot Creek, southern Flinders Ranges: (7), G), (1). Gm).
in), (0) Holotype $175; narrow, subcylindvical eolurins; (k)—S519; basal columns arising
from undulating stramatolite,
114
column margins, liminae wre completely re-
constituled to mw coarse, Xenotopic. polyganal
calcite Mosaic. grain size up to 0.5 mm; these,
in turn, contain subhedral dolomite crystals. as
well us disrupted rermnants of mieritie laminne.
The origin of the unlamingted selvage is pot
Clear, wherever it was observed, laminae are
somewhat coarsely recrystyllized immediately
adjacent to it inside the coluiin, and the sel-
vage may simply be the outermost lamina of
the wall preserved from rectystallization, bul
this is not certain since the selvage is unla-
minated. and laminae cunnol usually be traced
directly into it. There ure at Ieast two genera-
tions of calcite veins: the earlier ones are more
imegular, finer grained, and contain dolomite
rhombs, suggesting that they pre-date at least
one period of dolomitization. The younger
veins are straight, more coarsely crystalline.
and post-date dolomitization. Dolernitization
in these stromiatolites is, ut least in part, very
late Giugenetic.
Coniparisons
In having long, straight, infreyuently
branching columns without rapid changes in
Uiameter, these stromatolites are distinguished
from idl but Minjariae Krylov and Surusania
Krylov, They are distinguished from other a-
or #-parsilel branching stromatolites (Boxenia
Korolyuk, deaciellt Walter and Katavia Kry-
lov) by their infrequency of branching. Min-
Jatia Krylov, however, has a ubiquitaus wall
und. lacks pedks and overhanging laminae,
Jurusaunia Keyloy may have either a patchy
Wall or no wall, pumcrous peaks, und [re-
quently a selvage covering columns, /. burren-
sis. is intermediate between Minjaria und Juru-
série Hut ass wssizmed tu the latter because of
its patchy wall and the presence of peaks...
hurrensis dillers Tram J. evlindrica Kryloy in
having a better developed wall, smaller and
fewer peaks, and less well defined lamination;
however. lamina shape is similar. J. deme
dunia Krylov is distinguished by its consistent,
well defined ribs and general ubsence of a wall,
J. burrensis 15 distinguished from J. nisvensiy
Raahen by its. inuch more even, smooth
columns which do not grade into or uhernate
with ps¢udocolumns and juterally linked
stromatolites; also, there ure no sharp changes
in limina shape as in the latter form, J, jude-
miei Kamar & Semikhatoy hus targer, often
strongly elongated columns, lacking a wall. J
derbalenis Bertrand-Sarfuti und J. alta Bers
tranw-Sorfari vlso lack wulls and have ragged
column ntargins, J, lisse Berttand-Sarfau is
W. V_ PRETSS
distinguished by the absence of peaks and cer-
niccs, and by more frequent branching,
Distributions Upper limestone band ot
Wundowic Limestone Member, Burr Well,
northern Flinders Ranges, S. Aust.
vtge; Late Adelaidean, correlyted with the
Late Riphean or Vendian of the USSR.
Group KATAVIA Kryloy
Karevir Krylay 1963: 94. Rauben 1969 83.
Glaessner, Pretss & Walter 1969: 1057.
Type Form: Katatia karatavica Krylov.
from the Katuv Suite of the Southern Urals.
Diagrens: Predominantly @-parallel branching
straight. subcylindrical, walled columns with a
markedly bumpy murgin structure.
Centent; Katavia keretaviea Krylov und
Katavia costata 1. nov.
Age: Late Riphean.
Katavia costata (, nov.
Katavia sp. nev.
Walter 1969; 1057.
FIGS. 6-0. 9g, (Oe Lob-d, 17u
Marerial: Seven specimens from near Depot
Creck, S. Aust.
Holotype: SUTS (Pips, 61, 1, myn. 0. beat)
from the Brighton Lintestone equivalent,
Depot Creek, southern Flinders Ranges. $.
Aust.
Nene: Latin costata, meaning “ribbed”,
refers (o the short ribs. present on the lateral
surface of columns.
Diagnosis; Katavia with very closely spaced
parallel columns, a thin wall, very indistiner
and wrinkled Jaminae. and a prominently
bumpy and ribbed margin structure with some
very short pointed projections,
Descriprion
Mode of Occorrenee: The stromatolites form
two Jenticular bioherms, 5m thick, and up te
100 m long. in the upper, pink dolomite mem-
ber of the Brighton Limestone equivalent. The
basal one metre consists of wrinkly flatelamina-
ted dolomite, with concordant atylolites. This
zone gives rise directly Ww narrow, parallel
columns (Fig, 6k), which continue through-
out the height of the bioherm (Pig. 16b). At
Frequent intervals columms cut by horivantsl,
concordant stylolites, The upper surfaces of
hioherms not exposed. At margins of bidherms
columns became inchned at »hout 45° (Fig.
17a), hut no horizontal columns were observe.
Colum Shape ane Acrungenent; Columns
long, straight, yerv closely spaced, diam. 05-3
Glaessner, Preiss &
SOUTH AUSTRALIAN STROMATOLITES I
cin, most commaniy 1-2 em (Fig. 16d). Most
columns vertical, except near bioherm mar-
gins, Cross-sectians round to polygonal, often
resembling iid-cracked polygons (Fig. 16c}.
Columns may be 5-20 em long between
branches: occasional columms only a few cm
long have pointed terminations (Figs, 61 J. 1.
m. n, ).
Branching moderately frequent, predominantly
-parallels a column 1.0-1,5 cm diam. widens
gradually to 2-3 em, then divides into two, less
often three, aurrower columns (1-1.5 ¢m in
diameter}, u-parillel branching from broad
colurnns does nat occur, Sone branching very
slightly divergent,
Murvitr Streedure: Lateral suctace of colunins
markedly bumpy and ribbed (Fig. 61-0}.
Fquidimensional bumps. 0.3-1,0 cent diam,,
with a relief of 2-5 mm most common, These
prade into |runsversely elongated ribs, which
partly surround the columns, Small. pointed
projections up ta | em long moderately fre-
quent (Pig. 6i, 1). and in places slight niches
in the column margin (Tig. 16d), Overhanging
peaks extremely rare; bridges absent in speci-
mens studied Near column nvrgin, laminae
4urn down steeply to cover lateral surface for
short distances, so that onty two or three la-
minge form the wall (Fig. 16d), which is deve-
laped almost everywhere, cuveriag all bumps,
Tibs jind projections.
Liwitece Shaper Laminge in basal, Mat-lamina-
jed portion poorly preserved, but appear Lo be
wivy and wrinkled. The lowest narrow
columns venerylly Have gently convex, wavy
and wrinkled lalinae, but degree of convesily
increases upwards, Undulations have wave-
lengih 2-5 1m. Fig. %¢ illustrates commonly
occuring famina shapes, 62% of laminae
measured have b/d hetween 0.3 und OS (Fig.
102), Most laminae hemispherical, some wp-
proach rectangular shape. Luminae near the
most hurpy column margins commonly
strongly wavy,
Micrastructere: Lamination in all specimens
extremely indistinct. Where best preserved, it
consists. of alternating relatively lighter and
darker, pale brownish stained dolomite Ja-
mings. many of the light laminae containing
uctrital quartz sand grains, restricted to the
central paris of columns. Liehd laminae have
extremely indistinct boundaries, are 0.3-2.0
mm thick, and thin myrkedly towards column
margins. Included sand grains subrounded to
subanvular, yrain size 0.05-0.5 om. Balomite
hypidiotopic, of inequidimensivnul erystals.
Wl
grain size 0.005-0,025 mm, often showing up-
proximate rhombic outlines, There are vatin-
tions in the intensity of the brownish pigmen-
Igtion present In the crystals, Dark lamilnee
extremely fine grained, more densely stained
teddish-hrown, O.05-0.5 mtn thick, most
clearly visible and thickest in marginal por-
lions of columns, but thin, markedly wrinkled,
unc discomlinucus, frequently consisting of
lenses only | or 2 mm long, in central part.
Towurds margin, dark laminae frequently
merge
Mnterspacey extremely niyrrow, 1-3 mm wide,
most commonly 1—2 mm. Sedinventury Alling
Unliminaied, comasis of equal propurtinns. of
sand and dolomite nratrix, Quartz sand grains
subrounded, cummonly 0,2-0.5 mm diam. a
few up to 2 mm. Feldspar and red. extremely
fine vrained, possibly igneous rock CLragments
sub-ordinatc, Matrix consists of hypidiotopic to
xenotopic dolomite, with equidimensional crys
tuls of grain size 0,005—0.03 mm, patchily re-
erystallized ta hypidtotepic sparry dolonite of
0.03.05 mm grain size. Intraclasts of pale
brownish fine grained dolomite, up ta 5 mm
fang. 2 mm wide occur in places mixed with
sand jrains. These probably represent frag-
mented alval laminie.
Secondary Alteration; The generally poorly
preserved microstructure of stromalolitic and
interspuce dolomite and its corrosion of quurlz
Brains suggest that it is secondary. Small irre
gular patches of recrystallized, fine sparry dolo-
mite arc scattered throughoul columns ans
interspaces. Layering in stramatolitic columns
is extremely tndistinct, and defined only by
slight vanations in grain size and pigmentation;
this general homogeneity may be partly duc to
dolomitization. Dark laminae are in places dis-
fupled, perhaps by recrystallization of the
intervening light laminae. All detrital quartz
grains have corroded margins, nsually sur
rounded by a thin tim of Jinely crystalline
spurry colomite. Authigenic chlorite is deve-
loped in places in the interspace sediment near
enlumo margins. Small stylolites are developed
locally near column margins bud are unimpor
lant, Prequent (urge stylolites, concordant with
bedding, were seen in the field (Fig. 173),
These we up ia 1 om wide. ond contain
marked concentrations of sand and auchiuenic
chlorite,
Comparisons
The stromatolites are assigned Lo the group
Ketavia becuuse of their A-parallel branching,
bampy, walled columns. They ine distinguished
92 W. V. PREISS
etl
5cm
—e
3.4 Depotment of Hines
Fig 7. (4) to (m)—Kulparia kulpurensiy, Etina Formation equivalent, near Kulpara, northern
Yorke Peninsula: (a). (6b), (c), (e), (i) —Holotype $380, frony unit C (Fig. 8}; (d), (m)
—S419; junctions between two contiguous domes, unit C (Fig. 8), (m) is cut by a sand
dyke, including stromatolitic fragments; (f), (g¢)—S420, from unit E (Fig. §)3 (h)—S271,
from unit C (Fig. &); (j)—S381, (k), ()—S270. from unit A (Fig. 8).
SOUTH AUSTRALIAN STROMATOLITES II i
from most other walled stromatolites by their
markedly bunypy murgin structure, and from
Patomla Krylov by their predominant simple,
B-parallel branching. Like the illustrations of
Katavia karatavica Krylov, K. cestata has. a
few very short pointed projections. It is ex-
tremely similar wo K, Acratavicu in tts gruss
form, micrustructure and margin structure, and
is distinguished only by ils. more closely spaced
columns and by the possession of short trans-
verse ribs,
Distribution: In two iioherms. upper (dolo-
mite) member of the Brighton Limestone
equivalent, 3 km N of Bepor Plat HL.S.,
southern Flinders Ranges, S. Aust.
Age; Lute Adeljidean, correlated with the
Late Ripheun of the USSR-
Group KULPARIA Preiss & Waller
(in Wulter 1972: 151)
Pulomia sp. nov., Glaessner, Preiss & Wal-
ter (1969, p, 1057).
Type Form: Kulparta hulparensis Preiss,
from the Etina Formation equivalent, Wm-
heratana Group; Yorke Peninsula. S. Aust.
Name: After the township of Kulpara,
northern Yorke Peninsula, &. Aust.
Diagnosis: Long, nearly straight, parallel
bumpy columns, erect or radially arranged
with very frequent coalescing and bridging,
moderately frequent u- and A-parallel branch-
ing and a wall between bridges: projections
may be moderately frequent,
Canrent: K. kulparensis Preiss and KR. aliete
(Cloud & Semikhatoy) Walter.
Comparisons
In gross form, Kidparia resembles Minjaria
Krylov und Boxenia Korolyuk, but is distin-
guished by its bumpy column margins with Fre-
quent bridging and ecovlescing, Like Natavia
Krylov and Paonia Keylov.. it has a walled,
bumpy margin structure; Katavia colinns
have /-parallel branching, no bridges and they
rarely coalesce, while Patontia hus frequen
slightly divergent branching and very nu-
merous pointed projections, Some illustrations
of Palomia ossice Krylov. from the Malokaroy
Suite, resemble AKudparia in having bunypy,
long subpurallel columns with fewer projec-
tiuns, but lack the frequent coulescing und
delicate bridges of Kulperia. Kulparia kul-
parensis was initially assigned to Patamia on
the busis of this similarity (Glaessner, Preiss
& Walter 1969). Kuloaria differs from Linefle
Krylov in lacking gnarled and tberous
columns, and trom Gywnesolen Steinmann in
lacking y-parallel branching. in gross forfn,
Rulperla alsa resembles the walled parts of
Inzeria intia Walter but is distinguished by the
absence of niches and elongated projections,
Distribution; Elina Formation equivalent,
S. Aust. and Bitter Springs Formation, €
Aust,
Ape! Adelaidean.
Kulparia kulparensis f. nov.
FIGS. 7, 8, 9h, LOh, 16e, b7b-f
Material: Eleven specimens trom Kulpara.
S. Aust.
Holeioype: S380 | Figs. Ja, b,c. & 1. 17d, a)
from the EBtina Formation equivalent, Kul-
para,
Name: After the township of Kulpara,
Diagnoyis: Kudparia with very frequent delicate
bridges, moderately frequent pointed projec-
tions und variable lumina shape, from gently to
steeply convex. Microstructure diffuse, irregu-
arly streak v.
Descriplion
Mode of Occurrence: A bed traced for at least
400 an, its noarthern extension not Known,
while its termination in the south can be lo-
cated only approximately, due to lack of expo-
sun. Strramatolitic bed up to 13 m_ thick,
occurs at passage from flaggy pale grey clean
limestane lu massive, grilly. cross-bedded
limestones, The basal portion of the bed (A)
(Fig. 8), commencing contormably upon the
fiagey limestones, consists of short, partly
divergently branching columns and psctudo-
columns, in thin beds up to L5 om thick, with
numerous bridges and continuous, nearly fat-
laminated layers. This is overlain by a broadly
domed biostrome (C) of long, narrow, vertical,
parallel, very closely spaced columns, arising
from a laterally linked zone and short, browd,
basal columns (B). ‘The upper surface of the
biestrome of long parallel columns hends
downwards sharply at the junctions between
domes, columns becoming inclined, and io
some extent pseudocolumnar, The domed bio-
strome pattern is repeated im the overlying un-
dulose and pseudocolumnar bed (D), again
passing up into long, parallel columns (E),
once more overlain by laterally linked and
pscudocolumnar lJayers (F), Gritty, cross-
bedded Jimestone overlies the stromutolitic
seyuenee.. Contacts between the various units
cannot be accurately placed in the field, clive
tO poor exposure and lJichen cover, but were
Hg W. Vo PREISS
pardy deduced from jaboratory study of speci-
mens (Fig. 8).
Column Shape and Arrangement: Unit (A)
consists at short, yertical to slightly inclined
columns, 5-20 mm wide branching frequently
from u wavy laminated layer. Columns swell
and constrict slightly, bear rounded bumps and
vecusional ribs, and coglesce frequently. Some
columns terminate their growth as pointed pro-
jections (Figs, 7), Kk. 1 978); overlying unit
(B) in part. columnar Tf present, columns
broad, up to 6 em wide, with very irregular,
bumpy outlines and numerous massive bridges.
grading laterally ond vertically into pseudo-
columns. with uvecusional interspaces, In the
main columnar units (C) and (E), columns
1-3 cm wide, swelling and constricting slightly
(Big, 17d. c). A few branches develop only
inta short, puinted projections (Fig. 7f, g).
Length of long, parallel columns between
Brunches 5-2(h.cm; the unit asa whole attuins
a thickness of up to 2m. but columns. not con-
linuous thrsughout, as pseudocolumnar hori-
zons intervene. ‘I famsverse
rounded polygonal, lobate. elonguted or irre-
gular: circular sections relatively rare, At dome:
edves, colunins slightly inclined (never at less
thun 607 to the horizontal), and bridged to
a greater extent, forming pseudocolumns
resembling those in units (B) and (D) (Fiz.
7d. im)
vi iat ow nf
Nadeau 1 L,
= eweneel BAA iat
jal PAVE YE 4 SY
‘ haces Fisas¥s ay rae
Soy es t+ Et
pb io!
Voncbir bide
Sy Deyatenent ef snes
Fig. § Disgrimmatic section of stromatolitic
bed near Kulpara. The felalive positions
of the specimens weie partly determined
in the field aml partly dednced from
laharntary specimens,
sections generally’
Branching: Basal columns of unit (A) chacac-
tenzed by frequent, slightly divergent branch-
ing (Fig 7}, k. 1). Long, parallel columns of
units (C) and (8) entirely a and A-parallel
branching, Near their biases, broad columns
and pseudocolumns (4-6 cm wide) hreneh
into several I—3 em columns. Above this level,
az and @-parallel branching moderately fre-
quent. Coalescing of neighbouring columns is
as frequent us branching.
Margin Siructire: All columus have a mark-
edly bumpy literal surface; bumps 0,5—1.0 cm
wide, with a relief of 1-5 mm most com-
mon, Most equidimensional, some grade into
short transverse ribs, others into short pointed
projections (He; W, 2), Longer pointed projec-
tions (up do 3 cm) moderately rare (Pig. 72)
Delicite bridges, compased of only one or twa
laminac very frequent, linking most adjacent
columos (Pig, 17d), usually depressed, U-
shaped (only the more prominent bridees
could be shown on reconstructions). Massive
bridges up to 2 ¢m thick moderately rare. Suc-
cessive delicate bridges in places only 5 mm
apart. Oecasionally very short peaks project
down from the column margins. Wherever
peaks and bridges do not occur. wall well deve-
loped. Wall most extensive in the lang, nurrow
columns. Laminae thin towards margin, and
coat suffuce for a distance of tip to | 5 em.
The wall involves from one to five laminae
(Fig. 17d, ©). ‘Vhe short basal columns of unit
(A) have only a patehy wall, as do some of
tke Jong columns with gently convex laminue
(Fig. 17b).
Lamina Shape very variable; generally narrow-
est columns have steepest laminae. while browd
basal columns and pseudocolumns have gently
convex and rectangular laminae. OF the
laminae measured, 695% have ratios of h/d
between 0.3 and 0.8, bul narrow columns and
projecnons usually have h/d greater than 1.0
(Fig. 10h). Fig. 9h illustrates coniinonty oeeur-
ting lamina shapes, Most laminac gently wavy.
usually with wavelength 2-3 mm.
Microsrruenire: Lamination indistinet «and
streaky (Eig, 17d, e), Where best preserved.
fairly continuous wavy dark liminae persist
from wall to wall, and alternate with light
tuminae. Dark laminae composed of very fine
grained silty limestone, consisting of equi-
dimensional xenatopic calcite of prain size
0.003-0.01 mm. with included subrounded
quarté and a little feldspar, ol grain size up
100.08 ran. Dark laminae 0.05—0.4 mim thick,
SOUTH AUSIRALIAN STROMATOLITES II
SA Depecttient oF Mines
. Examples of lamina shapes of the stramatolites, traced fram thin sections. (a)—Camephyton
garganicum garganicum: (b)—Gymnasolen cl. ramsuyi, (c)—lasgeria cf. tyonrest; (d)—in-
zeria conjuncta, (e)—Inzeriq: multiplex, (£)—Jurusania burrensix, (g)—Katavia cosratay (bh)
—Kulparia kulparensts.
generally thickest in central part of a column,
Boundaries’ diffuse. At column margins, dark
laminae thin to a thickness of about 0.05 mm
and coat surface of column. Intervening, light
laminue thinned more, and lens out some dis-
tance down the wall. so that here dark Jaminuc
merge. Light laminae up to 0,7 mm thick in
central parts of steeply convex laminated
columns. bul. thin rapidly towards the edges.
They consist of inequigranular xenotopic to
hypidiotopic calcite of grain size 0.015-0,05
mm, with minor rounded quartz. silt, of grain
size up to 0.08 mm. In the shert columns of
unit (A) lamination is better preserved (Fig.
{7f), Dark. homogeneous laminae, 0.15-1.0
mm thick, are composed of pale brownish and
greenish pigmented, almost equigrunuliar xeno-
topic calcite, of grain size 0,003-9.01 mm,
with inclusions of detrital quartz silt of grain
size 0.02-0.04 mm. Ih places, they have sharp
lower boundaries, but grade upwards into light
laminae, which are 0.3—1.5 mm thick, but thin
towards the column miurgins, and are com-
posed of slightly coarser, silty, xenotopic cal-
cite, of grain size 0.015-0.02 mm. Detrital
quartz grains are up to fine sand size (0.2
mm). All laminae extend uninterrupted across
the width of columns, unlike laminae in the
upper, long patallel cotumms.
Interspuces. wenerally very narrow {1-5 mm)
in units (C) and (EF), but wider in basal
columns of unit. (A). Their sedimentary’ fill
includes medium to coarse clastics, both terri-
genous und carbonate. Generally. quartz much
coarser than that incorporated inta columns,
The sediment consists. of approximately 40%
quartz (well-rounded, grain size 05-3 mm,
finer vrains tending to be subangular), 3%
feldspar {rounded to subangular cloudy micro-
NEI
MEAS RLD
Fig. 10, Frequency
SN, Deparemenc af Mires
distribution of lamina convexitics h/d for stromatolites iWlustrated in Fig. 9.
116
cline, up to 3mm gran size), 5% rack frag-
ments (rounded fragments. up to 4 mm, of
quartz-feldspar rock, quartvite and rare chert),
30% ¢arbonate allochems (inehiding fiat
pebbles 2-4 min Jong, flat pebbles coated with
3 or 4 pale and dark luminoe, recrystallized
oaids with dolomitic rims and rare composite
grains cemented by dark dolomitic rims) and
20% cement (sparry, hypidiotopic mosaic cul-
cle, of grain size O.015-0.06 mm cementing
allochems and lerrigenous detritus. in places
replacing the rims of these grains), Sediment
poorly bedded. Presence of wall between
bridges on columns indicates that sediment was
filled in periodically. Aller one influx of sedi-
ment, a bridge formed over it, then the inter=
space remained vacant while the column grew
another centimetre wor so. before the next
influx.
Secondary Alterwton; During diagenesis, the
carbonate of the long columns was partly re-
crystallized and dolomitized; some dark lami-
nue were preterentiglly dolomitized. and clays
were apparently redistributed inte a fine net
work of cracks and stylolites (Fig. (7e). In
places, the shape of laminae is completely dis-
rupted. Near the dome margins, lenticular
patches of spurry calcite occur within columns,
either concordant with the Juminae or at a
high angle to them. Vhese structures predate
clastic dykes which cut both stromatolile
columns and interspace sediment (both of
which must have been lithified at the time}
(Fig. 176). The filling af the dyke consists of
angular to subrounded, poorly sorted quartz, of
grain size 0.05-1 mm, The sand is tightly
WY:
PREISS
packed, the finest angular grains forming the
matrix. Calcite cement is almost totally absent}
quarl% grains are coated with iron oxide tims
In places, the filling process has actively eroded
the walls of dykes, so that disoriented trag-
ments of the surrounding Jimestone occur as
inclusions in the sand (Fig, (7b), Vhe dykes
probably formed by jointing of the alneady
lithified stromatolitic bed. especially between
adjacent domes, Concordunte stylolites, caneen-
trated at definite levels in the structures, Where
ihey are anly 1 or 2 mm apart, clearly post-
date the sand-dykes. Stylolites purtly follow
the tamination. and partly cut across i, Ver-
tical calcite veins up ta 1 cm wide, consisting
of course, euhedral crystals, post-date the stylo-
lites, and ate especially prominent in the junc-
lions between domes, which were persistently
subject t jointing. Dolomitization apparently
post-dates the formation of veins and stylolites,
und ts therefore yery late diugenctic.
Comparisons
These stromatolites have already been com-
pared to other groups. Ku/puria ktlparensis is
distinguished from K-. alieia (Cloud and Semi-
khatov) Walter, by its frequent delicate
hridges, generally more steeply convex laminae,
and by the presence of moderately frequent
pointed projections.
Distrihution; Etina Formation equivalent,
Umberatana Group, 7 km south of Kulpara,
northern Yorke Peninsula, ‘S$. Aust.
Age. Laie Adelaidean, correlated with the
Late Riphean or Vendian of the USSR-
Acknowledgements
T am indebted to Prof. M. + Glaessner for
supervising this study, to the support given
by the Centre for Precambrian Research, Uni-
versity of Adelaide, and to Dr, M, R, Walter
for discussions and collaboration, Drafting by
the Dralting Branch, Department of Mines,
und by my wife, i3 gratefully acknowledged,
This paper is published with the permission of
the Director of Mines.
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BERTRAND-SaRFATI, J. (1972), Paléoécologie ve
certains stromatolites en récifs des formdtions
du Précambrien supérieur du yvroupe d'Atar
(Mauritanie, Sahara Occidental): création
d'esptces nouvelles, Pakwoveogr,, Palaco-
climatol.. Pataceecol. U1, 33-63.
Croup. PL E.. & Seminmatrov, M, A. (1969),
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M. ®. (1969). —Precambriaun columnar stro.
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Komar, VY, A, (1966),—Stromatolity verktinedu-
hembriyskikh otlovhery severa stbirskoy
platformy | jkh stratigraticheskoe znachenie.
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posits of the North Siberian Platform and
(heir stratigraphic significance). Trudy geol.
Tayt. Leningr. 154, 1-122.
Komar, VY. A., Raaben, M. E., & SEMIKHATOV,
M. A. (1965).—Konofitony Rifeya SSSR i
ikh stratigraticheskoe znachenie. (Conophy-
tons of the Riphean of the USSR and their
stratigraphic significance). Trudy geol, Inst.
Leningr. 131, [-72.
Korovyuk, 1. K. (1963).—Stromatolity verkh-
nego dokembriya. (Stromatolites of the Upper
Precambrian.) /2 ‘“‘Stratigrafiyns SSSR” 2,
“Verkhniy Dokembriy”. (Moscow: Gosgcol-
tekhizdat.)
KryLov, 1. N. (1963)—Stolbchatye vetvya-
schchiesya stromatolity rifeyskikh otlozheniy
Yuzhnogo Urala} ikh znachenie dlya strati-
grafii verkhnegao dokembriya. (Columnar
branching stromatolites of the Riphean de-
pesits of the Southern Urals and their sig-
nificance for the stratigraphy of the Upper
Precambrian.) Trudy geol. Inst. Leningr. 69,
1-133.
Kryiov, I. N. (1967).—Rifeyskie i nizhnekem-
briyskie stromatolily Tyan’-Shanya 1 Karatau.
(Riphean and Lower Cambrian stromatolites
of Tien-Shan and Karatau.) Trudy geol. Inst.
Leningr. 171, 1-76.
MasLoy, V. P. (1937)—O rasprostranenii kar-
bonatnykh vodorosley v_ vostachnay Sibiri.
(On the distribution of calcareous algae in
Eastern Siberia.) Problemy Paléont. 2-3, 327-
342.
Preiss, W. V. (1972).—The systematics of South
Australian Precambrian and Cambrian
Stromatolites. Part I. Trans. R. Soc. 8. Aust.
96(2), 67-100,
RaaBen, M. E. (1964).—Stromatolily verkhnego
rifeya Polyudova Kryazha; ikh vertikalnoe
vaspredeleniec. (Stromatolites of the Upper
Riphean of the Polyudov Mountain and their
vertical distribution.) Byull. mosk. Obsheh.
Ispyt. Prir. Qtd. geal. 39 (vyp. 3), 86-109.
RAABEN, M. BG. (1969).—Stromatolity verkhnego
rifeya (Gimmnosolenidy). (Upper Riphean
stromatolites (Gymnosolenida).) Trudy. geol.
Inst. Leningr. 203, 1-100.
Rozanoy, A. Y., MIssarZHEVsKIy, V. V., VOL-
kova, N. A. Voronoya, L. G., KRYLOV,
I. N,, Ketter, B. M., Koro.yux, I. K.,
Lenpzion, K., MtkHWYyAK, R,, PyYKHOVA,
N. G., & Stporov, A. D. (1969) —Tommoat-
skiy yarus i problema nizhney granitsy kem-
briya. (Tommotian stage and the Cambrian
lower boundary problem.) Trady.. geol. [nst.,
Leningr. 206, 1-380.
SEMIKHATOV, M. A. (1962),—Rifey i nizhniy
Kembriy Yeniseyskogo Kryazha. (The
Riphean and Lower Cambrian of the Yenisei
Mountaing) Trudy. geol. Inst. Leningr; 68,
1-242,
Semikuatov, M. A., Komanr, V. A., & Sereprya-
Kov, 5. N. (1970)—Yudomskiy kampleks
stratolipicheskoy mestnosti. (The Yudomian
Complex. of the stratotypical area.) Akad.
Nauk. SSSR Trudy. 210, 1-207.
STEINMANN, G. (1911).—Uber Gymanosalen ram-
sayi, cine Coaelenterate von der Halbinsel
Kanin, Ball, Soc, geogr. Finland. 31, 18-23.
Wacter, M. R. (1972).—Stromatolites and the
biostratigraphy of the Australian Precambrian
and Cambrian, Palaeontology, Spec. Paper,
No. 11.
LIK
Fig. VI.
Fie. 72.
Fig. 15.
Fig, 14.
Fig, 15
Figy 16
Fig. 17,
W. V, PRETSS
Conophyton gargauea¢n gareqnicaum, from a vatt in the Paratoo Diapir: (a)—Near-axrul
section in outcrop of two adjacent columns illusirating murgin structure. The pen is 15 om
lung. Arraws indicate regular column margins; (b)—Transverse sections of ¢olimns in out-
trop. illustrating hoth circular and lanceolate shapes. The pen js 1S ¢m long; (c)—Qutcrop
section of broadly domed basal zoie, which gives rise upwards to conically laminated volumns.
(j—Thim section of crestal zone, brecciated perhaps hy compacuon of lithified Llaminne
Specimen No, S277; (¢)—Details of lamination. iustrating macrolaminae, detrital earbon-
ute granites (indicated by arrows), und swelling of some laminac. ‘Thin section. specimen
No. $214; (f}—Details of crestal zone illustrating contorted and thickened famination, Axial
lpnatludinal seclion. Thin section. specimen No, $932.
(al—Conaphyten yarganicum gargenicuny (Spec. No. $214). Longitudinal axial thin vectian
iMusirating erestal zone, laminstion, and deflexed margins of kiminge. The thick continues
bands are actually otacroalaminae. Natural ssizeé; (b), (c)—Gynmasolen of. ramsayi, From
boulders in a conglomerate in the ‘Vapley Hill Formation, near Wilson. Flinders Ranges:
(b)—-S388. Longitudinal thin section of verticul, branching cclumns, with iniraelust breccia
in iuterspaces, Natural size; (¢)—S387, Longitudinal thin section of joclined columns inter-
preted us derived from a bioherm murgin.
(uI—Gyenoselen cf. ramsevi from near Wilson. Longitudinal slab of regular walled
columns interpreted to be derived from a bioherm centre. Specimen No, S388; (b) to (¢)—
fuzevia ef. tomtsi, from the middle limestane of the Wundowie Limestone Member, Burr
Well, northern Flinders Ranges: (b)—Colomnar purtiun of a hioherm illustrating columns
with niche-projections. Note siylolites at base: (¢)—Domed basal part of bioherm with
continuous lamination; the upper columnar portion is separated by a stylolitic zone, Hammer
is 3 em Jong, (d)—Columnar zone overlying continuausly laminated basal purtion of bio-
herm. Note stylolitic zone ut pencil point. Pencil 17 cm long: (e)—Longitudinul thin section,
illustrating subeylindrical columns with altered margins and interspaces, gently convex to law
Makes anaes and a niche-projection, ‘Ehe basal part is iuensely cut by stylolites, Specimen
Oo, S452,
(a), (b)—Hizeria conjuncta, Brighton Limestone equivalent, Depot Creek, southern Flinders
Runes} (a)—Longitudingl thin scetlon of broad, basal coldains with niche-projection, The
laminae are allernaling dark, dolomitic, and light, calcitic, Holotype, $442; (b)—Longitudinul
thin section of inclined, tubcrous columns from bioherns margin. Specimen No. S4uds ic),
(chi—Jnzeria multiples, Brighton Limestone equivalent. west of Mount Remarkable, southern
Flinders Runges; (c¢)—Longiludinal thin seution of vertical columos, Natural size. Molotype,
S385; (d)—Longinidinal slab of same 4pecimen.
(2)—Inzeria mittipler, Brighton Limestany equivalent, east of VYednalue, southern Flinders
Ringes. Longitudinal thin seclion. Specimen No, 849; (b) to (e)—Jurusania brirrerists,
upper limestone of the Wundewie Limestone Member, Borer Well, ourthern tlinders
Kunges: (bJ)—Smooth eylindrical vertical columns near a bioherm margin. Hammer is 30 en
longa: (c)—Contiguous Spherieal bioherms; ()—T ongitudinal thin scetion iMustrating di-
chotomois s-parallel branebing bi cylindrical columns, Specimen No. S482: (e)—Lon-
gitudinal thin section of narrower cylindrical columns with slresky microstrocuire: Holotype
$543.
(a) Jernsdeiq barvensix. Longitudinal thin seetion of columns anising from undulating
slromatolites at buse. Specimen No, S481; (Bb) to (d)—Karavia cosata, from colomitic
‘inember" of the Brighton Limestone, Depot Creek: (b)—Long, vertical columns in longi
tidinal wulerop section. Hammer is 30 em lung. (¢J]—Transverse section of columns in out.
crop, Pon js 15 ¢m long; (d)—Longitudinal thin section iNustrating indistinet hirunation
and walled columns. Interspaces are lilled with sandy dolomite. Holotype, S!75. Natural
size. (¢)—Ardparia kalparensis, Etinn Formation equivalent, Kulpara. Parr of outcrop of
ovlindrical columns. Pen is 15 em Jong.
(u)}—Katavia costata, Brighton Limestone, Depot Creek, southern Flinders Ranges, Margin
of a bioherm showing inclinalion of columns ut right of photograph; (b) to (fF) Aulparia
ladparensis, Btina Formation equivalent, Kulpari. northern Yorke Peninsula; (b)—L.ongi-
tudinal thin section illustrating a sand dyke post-dating the lithification of the stromatolites.
Incorporated tn the dyke filling are fragments of the wall rock. Specimen No, 5420: (ce s—
Ouitrop transverse sections of lobute columns. Pen is 15 cm long, (d), (¢)—Longitudinal
(hin scetions of columns; holojype S380; (f)—Smoall irregular columns from Unit A at the
lose of the bed: longitudinal thin section. Specimen No. $270.
SOUTH AUSTRALIAN STROMATOLITES II
ia
FIG,
PREISS
W. V.
120
y2
FIG,
SOUTH AUSTRALIAN STROMATOLITES II
13
FIG.
122 W. VY. PREISS
SOUTH AUSTRALIAN STROMATOLITES II
W. V. PREISS
24
16
FIG.
SOUTH AUSTRALIAN STROMATOLITES II 125
STUDIES ON SOME SPECIES OF HAKEA (PROTEACEAE)
BY J. R. MACONOCHIE*
Summary
MACONOCHIE, J. R. (1973).-Studies on some species of Hakea (Proteaceae). Trans. R. Soc.
S. Aust. 97(2), 127-133, 31 May, 1973.
Hakea standleyensis sp. nov. is described from Standley Chasm, central Australia. This species is
allied to H. collina C. White of south-west Queensland but differs in leaf size and flexibility, and
fruit shape.
An examination of collections under H. multilineata has led to the recognition of five species:
H. multilineata Meisn., H. francisiana F. Muell. and H. grammatophylla (F. Muell.) F. Muell., and
two new species, H. minyma Maconochie and H. coriacea Maconochie. H. bucculenta Gardn.
which is allied to this group, is readily separated by its narrower uninerved leaves. The possible
evolutionary origins of the species in the group are discussed in relation to their distribution.
STUDIES ON SOME SPECIES OF //4KEA (PROTEACEAE)
by J. R. MaAconocie*
Summary
Maconocult, J. R. (1973).—Studics on some species of Makea (Proteaceae). Drans. RK. Sac,
S. duye 9702), 127-133, 31 May, 1973.
Hakew standleyensiy sp. nov, is deseribed from Standley Chasm, central Australia, This
species is allied to
flexibility, and fruit shape.
An examination of colleetions under H,
H. callinad ©. White of south-west Queensland but differs in leaf size and
muutilineata has led to the recognition of tive
species: A, multilineata Meisn., Hy franecisiana FP. Mucll and 2. grammatophylla OP. Muell.)
Fk. Muell., and lwo new species, HM. minveui Maconochie and M7. cariacea Mivonochie. Jf.
bueculenia Gardo., Which is allied to this group, is readily separated by its narrower uni-
nerved leaves. The possible evolutionary ovigins of the species in the group are discussed in
relation to their distribution.
1, Hakea standleyensis Maconochie sp. noy.
Frutex erectus diffusus, usque ad tom altus. Folia
linearia, teretin, erecta, infirme pungentia, circa
L5 mm diam. 30-65 mm longa (sed usitite
50-60 mm). Inflorescentia axillaris, racemosa,
parva, 6—-9-flora, rachide cu, 2 mm longa. Flores
ca. Bel] mm longi, per pilos longas sericeas albos
villosa, perianthio 4-6 mm longo atque pedicello
4-5 mm loneo. Ovarivm glabrum puene sessile,
ca, | mm longum, stylos glaber, ruber, ca. 9 mm
longus; stigma late conicum, ghabrum. Turns
obliquus, ghins rubra, semiannularis ad elongatum,
ca. | mm longa. Frveras ca. 15 mm longus, late
fulcalus, ca. 5 mm Intus Cad partem latissimam ),
pericurpo verruculoso, pedunculo ca. 10 mm
longo. Seminis corpus obovatum, ad apieem atten-
uatuim, cau. S mm longuni: ala ca, 5 mm longa,
reticulo tenui brunneo praedita,
Holorypus: D. J, Nelyon 1556, Standley
Chasm (23°41/S. 133°27'E). 53 km W of
Alice Springs, N.T. 19.ix.1967 (NT).
Iyvoiypis AD, BRI, NSW.
Specimens examined (all from Standley
Chasm). Chippendale & Johnsen (NT
3997), 16.x.1957 (AD, BRI, CANB. K.
MEL, NSW. NT, PERTH); Muconachie
464, 25.vili.[967 (NT); Nelyon 1555,
1Qin.1967 (MEL. NT): Must 356,
OX.AL9IOS (NT),
Erect straggling shrub up to Lom high.
Leavey linear. terete. weakly pungent-pointed,
erect on stems, about 1.5 mim in diam, and (30—)
50-60 (-65) mm long. Jnflorescence a srnall
axillary raceme of 6-9 flowers, the rachis
about 2 om long, villous with long white silky
hairs, Flowers about 8—LI mm long. perianth
4-5 mm long, pedicel 4S min long. Ovary
glabrous, almost sessile, about 1 mim long; style
glabrous, red, about 9 mm long; stigma
broadly conical, glabrous. Torus oblique:
gland red, semi-wnnular to clongate, about 1
mm long. Fra about 15 mm long, broadly
sickle-shaped, ubout 5 mm broad at the widest
point; pericarp verruculose, peduncle about
10 mm long. Seed-body obovate, tapering to
apex: about 5 mm Jong; wing about 5 mm
long with fine brown reticulations.
Habitat: on and in quartzitic rock ledges and
crevices almost at summit of maih outerop
ut reur of Standley Chasm.
This species is endemic to the Miedonnell
Ranges, central Australia, being found only
at high altitudes at Standley Chasm. Chippeo-
dale (1963) discussed the relic nature of plants
found in the Macdonnell Ranges und ft is pro-
buble that H. standlevensiy is a relic species,
possibly on the verge of extinction,
Hl. standlevensis is allied to H. colling C,
White (1944, p, 79) and also A. iiicracarpe
R, Br. H. collina is found on the sandstone
tableland of south-west Queensland and 7,
micrecaurpa is restricted to southern) highland
areas of castern Australia and extends down
to Tasmania. The three species may be separ-
ated as follows:
= Aril Zone Research Institute, N/T. Administration, Alice Springs, N.T. 5750.
128 J. R. MACONOCHIE
(1) Perianth glabrous. Leaves terete or trique-
quetrous a, coe. Hy microcarpa
(1) Perianth villous. Leaves terete.
(2) Leaves flexible, (30—)50-60(-65)mm
long, weakly pungent-pointed. Follicle
about 15 mm long, 4-5 mm wide, in-
curved along ventral edge . ;
FH, standleyensis
(2) Leaves rigid, 20-40 mm long, strongly
pungent-pointed, Follicle about 20 mm
long, 6-8 mm wide, almost straight
2 H. collina
along the ventral edge .
WERDARIUM. OF NE
ALICE GPRM
Fig. 1.
Il. Hakea multilineata and its allies
Bentham (1870) commented that he could
see no major difference between the descrip-
tions of H. multilineata Meisn. and H. gram-
matophylla (F, Muell.) F. Muell., except that
the raceme of the latter species has a densely
tomentose rachis, Bentham therefore con-
sidered H. grammatophylla as a variety of H.
multilineata, Black (1948) followed Bentham.
Bentham also placed H. francisiana F. Muell.
under H. multilineata although he did not see
HOLOTYPE
HEABARIUM OF MOATHERN TERRITORY
Eqvings, Auewabs
fe NT, ug
: ‘,
Ne Tae bien StS ye
(tip ae La, fee :
' Call Det
‘ Nates » ns ‘
Se pe
Holotype sheet of Hakea standleyensis Maconochie.
STUDIES ON SOME SPECIES OF HAKEA (PROTEACEAE)
any material of the first species. Eichler (1965)
treated A. francisiana as a separate species,
To study this problem further, collections of
this group of species were borrowed from the
principal Australian herbaria and also two type
sheels fram Kew and New York Botanic Gur-
dens, Data were compiled on inflorescence
length, colour and texture, the shape ond size
of fruit and seed, and lIcaf dimensions and
number of veins. The collections were initially
sub-divided into six groups based on gross simi-
larities and dissimilurities and then the mean
and standard deviation of the numbet of veins
per leaf (6 to 12 leaves per sheet) for each
group culculated, A t-test was then applied to
the data.
H. multilineatfa and its allied species have
the following similarities: all are shrubs; leaves
linear, flat with several to many nerves; infie-
rescence a many flowered raceme 2-I() cm
long, enclosed in bracts at the bud stage: stig-
matic cone long and narrow; fruit almost ses-
sile.
29
In general, the flowers of H. coriueeu, H,
francisiana, H. grammatophylla and H. multi
lineata are very similar. H. ntinyma differs by
having much smaller creamy-white flowers
(pink to red in the other specics),
H. hucculenta Gardn. (1936, p. 123) is
allied to this group of species in that it has a
raceme of similar size, shape and colour, simvi-
lar fruit. but differs in that its leaves are much
narrower und uni-nerved,
The mean number of veins per leaf, the
standard deviation and sample size are pre-
sented below—
Mean St, Dev. Sample size
A. frencisiana 6.0 1.3 182
H. grammatophyvila 7-1 1.4 84
H_ curtacea 10.3 5 163
ft. minyma 142 2.0 a9
AL omuledlineata 15.1 3.4 156
The probabilitics and t-values for difference
of means between species are presented in
Table 1.
TABLE 3}
Species 1
| A. cériaced 28.4 15.6 16.8 TWA
2 OH, francisiana ea 3.9 38.0 35.5
3H. grammataphylia <001 <,001- 27.0 21.6 tvalues
4 H.minyma <.001 << .001 << 001 25
5, ntultilineata <<ool <<00l <<001 .O1<P<.02
Probabilities
on ———————————
Thus at the 5% probability level, the sample
means of all species significantly differ from
each other. When the number of veins per leaf
is used in combination with inflorescence
length and size, pubescence on rhachis, und
fruit shape, then these species cai be readily
separated,
Key 10 Hakea multilineata aad allies
| Leaves with one distinct central vein
i, bucerlgata
1, Leaves with several to many veins -— — - 2
2, Muatuce fruit with distinet bicarinate ventral
suture Ssnbie d) H. niultilineqia
2. Mature fr wit “without bicarinate ventral
suture. vith fo bie eseterorsaryel np
3. Perianth creamy white _H. minyma
3. Perianth pink ta red. _... satlecsleeapitltrdte EE
4. Rhachis of infloreseence ‘tamentosc a.
HT. prammatophylta
4, Rhachis of inflorescence glabrous .___ 5
5. Main veins of leaves 5 or 7 (-8). Leaves 3-6
mm wide ....., ... A, francisiana
§. Main veins (8) 9-10. (- 13). Leaves 6-16 mm
wide ...... A. eoriacea
Hakea bucculernita Gardner 1936: 123.
Holotype: Gardner 2571 (PERTH) (n.v.)
Distributian: Restricted to Western Australia
from Galena in the north to Geraldton in the
south (Fig. 4).
Selected Specimens: W. Aust., Blackall 4709,
48 km N of Galena, 18.ix.1951 (PERTH);
Ginins 1550, 51 km S of Wartoo road house,
north of Geraldton, Aug. 1967 (PERTH):
Long 25, 129 km E of Geraldton, I.vii- 1960
(PERTH).
Hakea multilineata Meisn. 1847: 261,
Holotype; Drummond coll, fll no. 275, Swan
River (NY).
Tsotvpes: K (twa sheets);
MEL 1010216; PERTH.
Distribution: Restricted to an area in
MEL 10102132,
the
bau
south-west cotner of Western Australia (Fig,
5),
Selected Sperimens; W. Aust., Brooker 1872,
66 km EB of Brookton, 22.vii. 1969 (PERTH);
Drummond 275, Swan River (Vype) (K,
MEL, NY, PERTH); Filsen 8903. Holland's
Track, 88 km SW of Coolgartlie, 14.19. 196h
(MEL{, Wily 3220. ca, 120 km W of
Diniell on road wo Lake King, 15ix. 1964
(AD),
Hakea prammatophylla (F. Muell,) F. Mucll.
léh7- Zi.
Bayionyin, Grevillea grammatopliyla P. Muell.
1865; 24.
Holutype: “tn Australia centrali prope ecatral
Mount Stuart fructicibus intecspersa”, J. Wred.
Staart (MBL 1010236),
Mueller (1867) cited RK, T. Sullivan—-
“Guwler Ranges” and M, Weidenbach, “in
vicinity of Port Lincoln”, as 72, graumrate-
pivile but these two specimens are A. fronci-
siava FP. Muetl. The tragmentary nature of the
specimens probably explains the misidentifica-
tion.
Distribution, Restricted to the
central Australia (N.T.) (Fig. 5).
Selected Specimens: NLT. Beauglehole 23189,
King’s Canyon, George Gill Range. Svil. 1967
(NT, NSW)> Lothian 76, Standley Chasm,
Juty-Aug. 1954 (AD); Macorechie 443, Ser
pentire Gorge, 19.vil, 1967 (NT),
Makew minyma Macoiiochic, sp, nov,
Feutex L-2 m altus, caulibus mojoribos nonnullis
Peaeditus. Folia erecta, linearia, elongata, plana,
fuevia, glabra, rigide corincea, 8-15 em longa, 5-8
mm tata, o [4-17 nervis. (usitale 15) lineata, si-
floreseentia racemosu multiflora, thuchide glabra,
4-5 om fonga. Flores. maturi et expansi 8-9 mm
Jong. Perianthinm gilvam, glabrum: torus circa
| mm langim, OS ma tatum, horizontatls vel
aliquantum obliquus, Ovarium pane sessile, gluh-
ram, 1-15 nim longum; stylus glaber, filiformus,
3-6 mm longus; stigma labrum, ercetum, coni-
cum, 1 mo longum, Glans ovoidee—globosa, ad
basin avarii sita, Frets ovoideo—globusus, 2—
25 eny Jongus, 1-1.5 cm Jatuy: pedicellus 1-3 mm
longus vel minusculus; rostrum perconspicuum,
saepe curvatuin, Pericarpus laevis nisi pustilac
Parvae, plus minus straminicolor. Semiuls serpus
8 mm longum; ala 1.? em Jonga, nigra. rhombi-
formis vel angilato-ovata, secus corpus imilatera-
liter, decurrens-
Holotvpus: Macanochie 846, about 84 km
W of Musgrave Park Station, S. Aust. (26°
20'S; 130°30’B), 30.ix. 1969 (NT). Speci-
men with flowers, fruits and photograph.
iwaiypie AD, BRI, CANB, K, MEL, NSW
MERTH,
ranges of
J. R. MACONOCHIE.
HM. microneura C. A. Gardner= nomen. inva-
lidum in Fairall (1970),
Strub (—2 im tall, with several main ytems-
Leaves crect, flat, tinear, elongate, smooth,
glabrous, rigidly coriaceous, 8-15 em long,
5-8 mm wide, with [4-17 nerves (mostly 15),
Inflorescence a raceme with numerous flowers,
rhachis glabroos 3.0-5.0 em long. Open mature
flowers 8-9 mm long. Perianth creuamy-yellow,
glabrous, torus about | mm long, 0.5 mm
broad, horizontal ta slightly oblique. Ovary al-
most sessile, glabrous, 1-1.5 mim long: style
glabrous, filitorm, 5-6 mm long; stigma pliab-
rous, erect, conical, | mm long. Gland ovoid-
globular, at base of ovary. Fruit ovoid-globu-
lar, 2-25 em long. 1-1.5 cm broad; pedicel
[—3 mn or Jess; beak strongly developed, often
curved, Wall smooth with small pustules, col-
oured beige to light tan, the Iutler colour often
more pronounced on beak. The beak is often
lost From fruit older than twelve months and
the wall becomes grey in colour, Fruit thea
1.9-2.0 em Jong and |.4-1.6 em brood. Seed
bedy & mm long; wing 1,7 cm long. black,
rhombic or angulato-ovate in shape, cecurrent
along one side of the body.
The specific epithet ts derived from the Pil-
jantjatjara word minyma (woman). an allusion
to the fruit's resemblance lo a woman’s breast.
Aistribution; This species extends from. ihe
Musgrave-Mann-Petermann Range complex
of South Australia and the Northern Terri-
tory down to the Tammin-Merredin urea in
the south-west of Westeta Australia (Fig, 5).
Seleered Specimens: ST. Duntop 2010, 48 km
NE of Mt, Davics Camp, Mann Runge, 31x.
1970 (AD. CANB, NT); Larz 941, ca. 129 km
NE of Mt. Davies Camp, edge of Pottoyi
Hills, N.T_, 2,xi. 1970 (DNA, MEL, NT), S
Aust, Eichler 17285, between Mi, Harriet and
Musgrave Park Homestead. S.ix. 1963 (AID):
Macererkie 846, ca, 84 kin W of Musgrave
Park Station, 30,ix, 1969 (Type) «NTI, W.
Aust. Gardner 839, Coolgardie, 4.x. 1920
(PERTH); George 2879. 35 km NE of Luver-
ton. 23.viil, L961 (PERTH); Gearge 5639, 55
km SW of Wiluna, 29%.vii, 1963 (PERTH):
Koch 975, Cowcowing, Sept 1904 (MBL,
NSW, PERTH): Rovee 4461, Comet Yale,
23.ix. 1953 (PERTH).
Makes francisiana F. Muell, 1858; 20,
Type: G. Francis, near bay, Spencer's Gulf.
Specimen probably lost (search made at AD,
K, MEL).
Nearype: RB. Copley 2345, Thirlga Station,
Cluwler Ringes, S, Aust, |3.x, 1968 TAD),
STUDIES ON SOME SPECIES OF HAKEA (PROTEACEAE) 131
i re
ee
as
Shy emadeg-
ABOLIMHSL NYIHIHON 40 Wh”
fee
cae, FeO
WALOIOW
O yn
ivis
Woeveeay
Fig. 2 (above). Holotype sheet of Hakea minyma Maconochie.
Fig. 3 (below). Holotype sheet of Hakea coriacea Maconochie.
{72 ]
tl. anultilinedta vir gramined
lidum in Bairall (1970),
Disiribution: Widely distributed theougl the
southern arid areas of South and Western Ausa-
tralia (Pig. 4),
Selected Specimeny: S. Ausl Cornwall 56, ca.
SS km SE of Kimba. ivi. 1968 (AD. NV):
Rimyay $.n,. 113 km SSW of Camp 17. Elder
nomen iWyVva-
Expedition, July 1891 (AD, NSW): Wilyon
(573, 40 kor NW of Ceduna. las. 1960
(AD). W. Aust) Gardner 6465, Bencubhin,
T0.ix. 1942 (PERTH): Georve S646, 122 kin
™ of Sandstone, 29.vil, 1963 (NSW, PERTH):
Hilyvan 31442 ca. 30 km SE of Londonderry.
Idix. 1964 (AD, PERTH)
&
sa
Distribution of A. fhuceutenta (m), H
coriuced (&), and Jf: franeisinne (®),
bin. 4.
RSs
hig 5.) Distribution of //. granunatophylla (®),
Ho ominyma (@). He miutrilineata (a).
From Kangaroo L, South Australia, there
huve been two sterile collections made of a
species close to /7. franeisiana, bur until better
material is availuble, its status is obscure,
Hakea coriacea Maconochie sp, nov.
Frifex 3-4 1 altus, Folia linearia elongata, plana.
eoriace’s intervenium pobescenti, 920 em Tonga.
usitiite [4-17 em. 6-16 mm lata usitate 8-10 mm,
» RK, MACONOUCHIE
s—l4 hervis interns plerumque 9 vel LOL Lafleres-
centia racemosa multiflora rose vel earneéa, cha.
chide glibra, @=! 1 em longa. Ploy glaber 23-25 mim
longus, pedicelluy 2-3) mm longus, perianthium
7-8 mm, stylus ghaber |9% 27 mm Jongus. stigma
Wabraum. conicum 1.5 mm longum. Torys ali-
qhanium cobliquus, ghins semi-annularis. Brae tea
elabra vel pubertila, caduea, marine ciliata.
Fractis ovoidea-globasus, circiler |& mnt jongus,
12 inm Jatus ef evassus, pericarpiis laevis nisi pus-
lulne putteae purvae vel aliquando fissuris paucts.
Affinis Haekede franeistinae Fo Muell sed differt
numero majoro nervorum et foliibus latrioribus.
Shrub to 3 to 4 m high. /eavey linear, Mat.
coriaceous, with a fine pubescence on the inter-
veinal area, 9-22 em long mostly 14-17 em.
6-16 mm wide. mostly 8-10 mm with S13
nerves. Inflorescence pink-red, rhachis glabrous
6-11 cm long, a raceme of many flowers.
Flower glabrous 23-25 mm long, pedicel 2-3
mm long, perianth 7-8 mm, style ghibrous
19-21 nin, stigmatic cone 1.5 mm long Terns
slighily oblique. ghind) semi-annular. Bracts
glabrous or sometimes puberulous with ciliate
margin, cuducous. Fruit woody. shortly pedun-
culite (2-3 mm) about 1S mm tong, 12 mm
wide and broad. wall smooth with w few small
pustules or sometimes with small fissures.
Closely related to AM. franciniane but differs in
greater number of nerves and wider leaves.
Holowpus: C. A. Gardier (2155, hetween
Perenjori und Jibberding, W. Aust. Sept.
1953 (PERTH).
Dis:ribmiien: Restricted to awn
WSW of W. Aust. (Pin, 4)-
Seleeied Speciinens: W. Aust. Aplin 1983.
3 kn EF of Tummin, 13.7%. 1962 (PERTH):
Drupiniond (8. W. Aust. (MEL. NSW);
Koch 1018. Coweowing, Sept, 1904 (AD,
MEL, NSW); Melville 4265, U8 km W of
Dalwaliina, 2l.vii 1953 (AD. BRI K.
MEL. PERTH).
ures in the
Phylogeny and Evolution
Tirese species form a natural group dillerine
from the other members of Benthan’s Hekee
scet. Conogvinoides ser. Longivivlee by the dis
distinctly elongate raceme, 2-10 em Jong. ‘The
other members of this series all have a more
compact raceme, resulting in a more globular
inflorescence.
The phylogenctic relationships of this group
are uneertain:
(1) The inflorescence. leaf and fruit structure
of HA. francisiena, HL. coriacea, MH. grammar.
phyvlla and I. bucealenra indicate they pro-
bubly have a common ancestor, and that J/
muiilineata and HH. tonvine may have evolved
independently,
STUDIES ON SOME SPECIES OF HAKEA (PROTEBACEAE) 133
(2) The similar distribution patterns of H.
frencixiane and H. minyma suggest that these
two. species may have had a common ancestor,
and //, coriacea, H. grammatophylla, H. buc-
culenta and possibly H. multilineata were all
derived trom Al. francivsiana.
The south-west province of Western Austra-
lia appears to be the focus of origin of this
group of species, as five of the six species
occur there and the distribution tends to
radiate from there into the more arid areas to
the north and cast.
The two records of Makeu ef. franecisiana for
Kangaroo Island suggest thal, during an ¢arher
geological period, Kangaroo Islund acted as a
migration bridge between Eyre and Yorke Pen-
insulas and Fleurieu Peninsula. Wood (1930)
refers to this connection and rewards it as
tecent in geological time.
The implication of these observations is that
either (1) this group of species may have
evolyed, diversified and migrated during the
period of a land connection between the Eyre
and Fleurieu Peninsulas or, (2) this was a
period of rapid spread of H. /ranersiana,
The restricted distribution of H. gramsiio-
phylla to the tanges of central Australia and
the distributional pattern of A. francisiana
would Indicate a north-eastern migration route
from the south-west province of Western Aus-
tralia, Subsequent periods of aridity would per-
mit speciation to occur as there was a retreat
to more favourable habitats.
Acknowledgements
The Directors and Curators of the follow-
ing Australian Herbaria (AD, ADW, BRI.
CANB, CBG, MEL, NSW, PERTH) are
thanked for allowing examination of their col-
lections and also Kew and the New York Bo-
lanic Gardens. for making available Drum-
mond’s type sheets,
To Mr. J, H, Willis, | am indebted for two
of the Latin deseriptions and to Dr. Hj, Eichler
for his advice on nomenclatural problems and
comments on the manuscript.
References
Benruam, G. (1870).—"Flora Australiensis™, Vel.
§. (Reéve: London.)
Brack, J, M. (1948).—Flora of South Australia’,
Part Il, 2nd edn. (Govt. Printer: Adelaide.)
CuHrpPpeNpALe, G. M. (1963).—The relic naire
of some central Australian plants, Trai, R.
Soe. 8. Asst. 86, 31-34.
Eicutrr, Hj. (1965).—*Supplement to J. M.
Black's Flora of South Australia”. (Govt.
Printer: Adelaide.)
Faimate, A. R. (1970).—“Western Australian
Nutive Plants in Cultivation’. (Pergamon:
Australia, )
Garpnern, C, A. (1936).—Contribuliones Florae
Australise Occidentalis No. IX, J. R. Soc. W.
Aust, 22, 123-128.
Meisner, C, F, (1848).—Jn ©, Lehmann,
“Plantae Preissianae” IJ, 260-262, (Hamburg.)
MUELLER, F. (1858) —“Fragmenta phytozraphisde
Australiae” Vol. LL (Govt. Printer: Mel+
bourne.)
MUELLER, F. (1865).—"“Fragmenta phytographiac
Australiae” Yol. Y. (Goyt, Printer: Mel-
bourne.)
MueELLeR, F, (1867)-—“Fragmenta phytographiae
Australiae” Vol. Vi. (Govt. Printer: Mel-
hourne.)
Wire, C. T. (1944).—Contributions fo the
Queenlsand Flora, No. 8. Proe. R, Soe. Qld
§5 (5), 79-80.
Woop, J. G. (1930),—An Analysis of the Vege-
tation of Kangaroo Island and the adjacent
Peninaulus. Trams. R. Soc. §. Aust, 54, 105-
139.
PHOSPHORIAN LAVENDULAN FROM DOME ROCK MINE,
SOUTH AUSTRALIA
BY A. W. KLEEMAN* AND A. R. MILNEST
Summary
KLEEMAN, A. W., & MILNES, A. R. (1973).- Phosphorian lavendulan from Dome Rock Mine,
South Australia. Trans. R. Soc. S. Aust. 97(2), 135-137, 31 May, 1973.
A new variety of the rare mineral lavendulan (= freirinite) has been found in Precambrian rocks in
South Australia. It is notable for containing a significant amount of phosphorous replacing arsenic.
Its formula, based on microprobe analysis, is (Naj.o7Ca1.91Cu4.go) (AS3.16P.s4)O16Cli ss 4H20. Powder
diffraction data are also recorded.
PHOSPHORIAN LAVENDULAN FROM DOME ROCK MINF,
SOUTH AUSTRALIA
by A. W. Kieeman* and A. R. MILNES}
Summary
KreeMan, A. W., & Micyes, A. R. (1973),.—Phosphorian lavendulan fram Dome Rock Mine,
South Australia. J'rans. KR. Soc, §. Ausf. 9742), 135-137, 31 May, 1973.
A new variety of the rare mineral lavendulan (= freirinite) has been found in Precam-
brian rocks in South Australia, It 1s notable for containing a significant amount of phos-
phorous replacing arsenic. Its formula, based on microprobe analysis, is (Nay .o7Car.o1Cuy.so)
CASy 43P.94)0i6Ch a5-4HeO. Powder diffraction data are also recorded,
Tntroduction
The Dome Rock mine is situated about 44
km ENE of Olary (148°27’R, 31°55/S). The
detailed geology of the mine area is described
by Dickinson (1942) and a brief description
is given by Campana & King (1958). The
country rocks are low grade metamorphics of
the Willyama Complex: Dickinson considers
the lodes ta be replacements of a fine grained
sandstone. The primary ore is reported by
Campana & King to be chalcopyrite and pyrite.
Some cobalt is found in the sulphides and ery-
thrite stuininys were reported by Mawson in
an unpublished report to the Dome Rock Cop-
per Mining Co, The fodes are oxidised to a
depth of about 60 metres. The oxidised ore was
inainly chalcocite, tenorite and cuprite, with
olivenite and chrysocolla.
Bayliss er al. (1966) examined specimens
of oxidised ore from Dome Rock and reported
several arsenic minerals occurring as “encrus-
tations alony partings in siliceous ironstone”.
They identified clinoclasite, Cu, AsO,(OH)..
conichalcite, CaCu(AsO,)(QH), and a third
mineral which they called chlorotile,
Cuy (AsO,)4.6H.O with the comment that it
also. =oresembled = mixite, Cu,,Bi(AsO,),;
(OH),,, 6H:sO. However, chemical tests
failed to reveul the presence of bismuth. They
also record the identification of cornwallite.
Cu, (AsO, ).(OH),.H,0 but do not quote
the authority, Their own identifications were
based on X-ray diffraction amplified by the
chemical test on the “chlorotile”.
Early in 1972, Mr, H_. Gallasch submitted
a sample from the 120 ft. level of Dome Rock
mine, containing a mineral which proved to
be quite different from any reported by Bay-
liss et al, The deseription of this mineral re-
sembles that of clinoclasite given by Buyliss
et al., but the powder patterns are dissimilar.
Tt occurs as rosettes of ucicular blue crystals on
u block of siliceous ironstone, The rosettes are
about 2-4 mm in diameter and the individual
crystallites are Jess than 0.02 mm_ across.
Broadening of the lines in X-ray powder phato-
graphs indicates that the mineral is in fact cx-
tremely fine grained. The powder X-ray photo-
graph suggested that it could be sampleite or
lavendulan (Guillemin 1956). Accordingly it
was decided to analyse it on the microprobe
ta confirm its identity.
Methods
A polished thin section of some fragments
af the mineral was examined in the C.S.1-R.0O.
Division of Soils’ “Geoscan” (Cambridge In-
struments) electron probe microanalyser, The
elements As, Cu, Co, Ca, Cl, P. Mg and Na
were detected during reconnaissance spectro-
meter scans, Chemical homogeneity of the
mineral fragments was checked by photo-
graphing X-ray scanning images of the ele-
ments of interest. Selected areas of the mineral
* Department of Geology and Mineralogy. University of Adelaide. Adelaide, 5. Aust. S001.
7+C,S.1.8.0, Division of Soils, Glen Osmond, S, Aust. 5064,
130
fragments were then analysed quantitatively
for these elements.
Under normal conditions of analysis!, stg-
nificant systematic drifts in count rate with
time were observed for all elementy. This effect
is possibly due to a combination of photo-
chemical degradation (McConnell 1969) and
thermal decomposition of the mineral under
the influence of the electron beam in the eva-
cuated specimen chamber (Sweatman & Long
1969). Loss of alkali elements from silicate
minerals with time under the influence of elec-
tron bombardment during microprobe analysis
is a well known but not fully understood
phenomenon (McConnell 1969; Siivola 1969),
and can be minimised by reducing the beam
current. In the present instance, however, the
count rate drift was. minimised by expanding
the electron bear so that it sampled a circular
area 50 microns in diameter at the specimen
TABLE i
Micruprobe analysis
1 2
AssO;, 36.4 44,8
cud 38,2 36.3
CoO 0.03 nil
CaQ 5.7 5.8
Cl 5.5 3,5
PO; 6.0 n.d,
MezO 0.04 od.
NasO 33 3.1
Total 95.2
O=Cl 1.2
Total 94.0
1. Lavendulan, Dome Rock Mine (analyst A. R.
Milnes)
2. Lavendulan, San Juan, Chile (Guillemin 1956)
TABLE 2
Stractural Formula Based an As-+ P—4
Na 1.070
Ca 1.014
Mg 0.010
Cu 4.800
Co 0.004
Total 6.898
AS 3.161
im 0.839
‘otal 4.000
of 1.553
A. W. KLEEMAN and A, R. MILNES
surface, and by driving the specimen beneath
the beam at 30 microns per second during
analysis, The accelerating voltage and beam
current were maintained at 20KV and 50nA
respectively.
TABLE 3
Comparison of the X-ray diffraction pattern of
the Dame Rock mineral with those of lavendulan
and sampleite
T.avendulan! Lavendulan?
Dome Rock San Juan, Chile Sampleite2
dA I dA I dA I
9,20 vs 9.77 100 9.60 100
7.403 w 7.0) 40 6.85 70
6.76 Ww -
4.98 WwW
4.83 Ww 4.87 50 4.73 40
4.60 Ss
4.37 § 4.4) 40 4.30 80
4.17 s 3.89 70
3.50 vw
3.38 vvw
3.24 vvWw 3.23 50
3.18 vVw
3.12 8 3.11 70 3.04 100
3.06 Ww
2.98 vw
2.92 m 2.90 20 2.89 50
2.74 vw 2.76 20 2.80 50
7.69 m 2.69 50
2.61 m
2.47 m 2.48 20 2.50 50
2.40 Vw
2.34 vw
2.24 vvw
2.10 vw
2,02 vw
1.95 w 1.97 20 1.91 50
1,90 w
1.84 9 vyw
1.81 vw 1.83 20 1,79 710
1.75 m 1.76 20
1.72 vw 171 80)
1.69 yyw
1,66 vw 1,61 50)
1.50) yw 1.55 20 gar 70
140 w 1.47 20 1.44
1.42 20 1.37 70
1.21 20 1,21 50
1. Diffraction data measured by J. G. Pickering
(C.$,1.R.0.) using a 19 cm camera and CoKa
radiation.
Diffraction data from A.S.1.M. cards Nos.
11-351 and 11-349,
"
1 Normal conditions of electron probe microanalysis
Accelerating valtage 20 kV
Beam current S0nA
Flow proportional counters
Counting time: 10.seconds (line and one background position)
Spectrometers: LiF crystal—CnKa, AsKs, CoKa, Caka
Mica crystal—Naka, PKa, ClKe, MgkKa
Beam fully focussed
PHOSPHORIAN LAVENDULAN FROM DOME ROCK MINE 137
The raw output data from the electron
probe were refined by the CDC 3200 computer
program MICANCOR (written by H. Rosser
of C.S.LR.O.), which incorporates the correc-
tion program MKRPRB6 (Oertcl 1971). The
mineral analysis reported in Table 1 is the
average of analyses of eight selected arcas of
the mineral fragments in the polished section,
Results
The X-ray diffraction pattern (Table 3) of
the unknown mineral (measured by J, G.
Pickering) is similar to the diffraction patterns
of the isostructural minerals Javendulan
Na(Cu,Ca) g({AsO,),;Cl4HsO and sampieite
Na(Cu,Ca) ,(PO,),Cl4-5H.O. Therefore a
structural formula was calculated on the basis
of a total of 4(As+P) atoms (Table 2).
The result given in Table 2 agrees quite
closely with the data given by Guillemin
(1956) except for the excess of Cl in our
specimen.
We have used the name lavendulan rather
thun freirinite in accordance with the list. of
New Mineral Names (Fleischer 1957).
References
Baviiss, P.. LawrRENCE, L. J., & Watson, N.
(1966).—Rare Copper arsenates from Dome
Rock, South Australia. Avst. J. Sci. 29(5),
145-146.
Campana, B., & Kine, D. (1958).—Regional Geo-
logy and Mineral Resources of the Olary
Province. Bull. geol. Surv. S. Aust. 34.
Dickinson. S. B, (1942).—The structural control
of ore deposition in some South Australian
Copper Fields, Bull. geol. Surv. S. Aust, 20.
FLeiscHer, M. (1957),—New Mineral Names.
Am. Miner. 42, 117-124.
GuiLtemin, C. (1956)—Contribution a la miné-
ralogie des arséniates, phosphates et vana-
dates de cuivre. 1, arséniates du cuivre. Bull.
Sac. franc. Minér. Crist. 79, 7-95.
McConneie. J. D. C. (1969),—Photachemical
degradation of a silicate in the beam of the
electron microscope. Phil. Mag. 20 (8th ser.),
1195-1202.
OsrTeL, A. C, (1971).—The calculation of re-
sults in __electron-probe microanalysis.
C.S.L.R.O, Division of Soils, Technical Paper
No. 9.
Strvota, J. (1969)—On the evaporation of some
alkali metals during electron probe analysis.
Bull. geal. Soc. Finland 41, 85-91,
SWEATMAN, T. R., & Lone, J. V. P. (1969)—
Quantitative electron probe microanalysis of
rock forming minerals. J. Petrology 10, 332-
379,
RHODACARIDAE (ACARI: MESOSTIGMATA) FROM NEAR ADELAIDE,
AUSTRALIA. Hl. ECOLOGY
BY D. C. LEE*
Summary
LEE, D. C. (1973).- Rhodacaridae (Acari: Mesostigmata) from near Adelaide, Australia. II.
Ecology. Trans. R. Soc. S. Aust. 97(2), 139-152, 31 May, 1973.
Serial collections of rhodacarid mites extracted by desiccating funnels from surface soil
(greatest depth: 4 cm), moss and plant litter, at two sites on the western slopes of Mount Lofty,
overlooking Adelaide, South Australia were studied, as were small collections of rhodacarids from
two sites on the Adelaide Plain.
The presence of two communities of hemiedaphic rhodacarid mites is demonstrated by differences
in the characteristic species of two sites and a significant association into two groups of the species
of one subfamily (Ologamasinae). Population density is higher in the wet, cool winter and where
there is substantial, decomposing plant litter. Variations are demonstrated between some species in
the number of generations per year, the time for occurrences of particular life-history stages and the
sex ratio. It is suggested that species of Athiasella prefer higher nutrient loamy soils, while
Gamasellus is almost confined to low nutrient, sandy soils.
RHODACARIDAE (ACARI : MESOSTIGMATA) FROM NEAR ADELAIDE,
AUSTRALIA. II, ECOLOGY
by D, C. Ler*
Summary
buc, BD. C. (1973).—Rhodacaridae (Acari ; Mesostigmata) from near Adelaide, Australia. 1
Keology. Trans, R. Soe. 8. Aust. 97{2), 139-152, 31 May, 1973,
Serial collections of thodacarid mites extracted by desiccating funnels from surface soil
(greatest depth: 4 cm), moss and plant litter, at two sites on the western slopes of Mount
Lofty, overlooking Adelaide, South Australia were studied, as were small collections of rhoda-
catids from two sites on the Adelaide Plain.
The presence of lwo communities of hemiedaphic rhodacarid mites is demonstrated by
differences in (he characteristic specias of two sites and a significant association into two groups
of the species of one subfamily (Ofogamasinae). Population density is higher in the wet, cool
winter and where there {s substantial, decomposing plant Jitter. Varjalions are demonstrated
between some species in the number of generations per year, the time for occurrences of
particular life-history stages and the sex ratio, It is suggested that species of Athiavell: prefer
higher nutrient loamy soils, while Gamascllus is almost confined to low nutrient, sandy soils,
Introduction
Rhodacarids are mainly predatory mites, and
are most common and diverse in form in
Southern Temperate regions. The present
work formed part of a study on rhodacarids
from the environs of Adelaide, South Australia
(Lee 1970!), Part L dealt with systematics (Lee
1973) and should be referred to for the
authority to names of rhadacarids collected.
Part IIT, dealing with. behaviour, is to be pub-
lished.
Most rhodacarids are hemiedaphic, being
free-living in surface soil, plant Jitter, or in
moss or other plants with a similar growth
form. Some taxa, however, are not hemie-
daphic. Thus, Hydreagama@sus, Litogeamusns,
Parasiliphis, Perisetus and Tangaroellus have
only been found in or near the littoral zone,
usually on rocky shares; Rhedacaropsis has
anly been found in the littoral zone on sandy
shores: Cyrtolaelaps or Eur)parasitus have
generally been collected from bird or mammal
nests or from but caves; Tangaroellus porosus
Luxton has usually been found under the cara-
paces of barnacles; and the two species of
Laelapronyssus have been found closely
associated with flies or termites. Ecological
studies demonstrating more limited habitat
preferences include only non-hemiedaphic
thodacarid mites. Thus. Aydregamasus littor-
dis (G. & R. Canestrini) mainly occurs in
tock crevices in a limited part of the littoral
zone(Glynne-Williams & Hobart 1952, Morton
1954). Rhodacarus and Rhedacarellus are
commoner in the deeper soil layers (i.e. they
are euiedaphic) and are limited to purts of
sampled areas (Sheals 1957, Davis 1963, Wood
1967a, Emberson 19682), Cyrraluelaps and
Euryparasitus are commoner in mammals’
nests that are on the ground and made of moss
(Mrciak, Daniel & Rosicky 1966).
There is a problem in defining precise habi-
tats for ground inhabiting mites, because a
species may occur in strictly limited habitats,
* South Australian Museum, Adelaide. S. Aust. 5000.
Yipr. D.C. (1970)—The taxonomy and general biology of the Rhodacaridac (Acari: Mesostig-
muta). M.Sc. thesis, University of Adelaide, Australia (unpublished).
“Emperson, R. M, (1968)— The Mesosiigmatu of certain coniferous forest soils in Western
Quebec, with a preliminary account of North American Rhoducaridae (Acarina), Ph.D, thesis,
McGill University. Montreal, Canada (unpublished).
144} Dp. Cc.
hut nt Widely differing localities. For instance,
it has heen shown in three different studies char
Rhodacarus roxens Oudemans has a limited dis-
tribution, It occurred only in a limited area jn
minecul suil over iron-stone in grassland ( Davis
19631; in non-calcaveous drift around wu lime-
stone Oulcrop in moorland (Wood 14¥67a}; and
in allvial saltmarsh (Luxton 1967), This type
of distribution for a number of species of mite
led both Davis (1963) and Wood ({967b) to
suxvest that many mite species consist of ecula-
gical races with different demands upon the
environment and therefore occurring in quite
different habitats.
The main aim of the present work was 16
demonstrate whether or not there ure any habi-
(at preferences amongst hemieduphic thoda-
carid mites. [nerdental information on seasonal
fNuctuations. of populations and fifc-histories
Was also sought, Th addition, becuuse species
association was considered in relation to halvitat
preferences, the correlation between taxunomic
affinity and degree of co-existence is discussed.
Four sites were initially sunpled befare two
were selected for serial sampling. The results
of the preliminary sampling from the two sites
that were not sampled again are also given, be
cause they suggest a possible correlution
WelWeen certain genera and environmental
factors.
Methods
Twe cxtraction methods were used: one for
dealing with disturbed “bag” samples collected
throughout a year and the other for undis-
turbed “core samples collected in August.
1. Bag Sampling and Extraction
Soil, down to a depth of approximately 4
cm, and the fitter or moss on it, was scouped
inte a plastic bag with u trowel. The volume of
a sample was about 1280 ml, and was taken
From in area of approximately 250 cm, This
sanple was poured intu an aluminium tube
(14% 40 em) with a wire mesh bottom, which
was placed on uy coarser wire mesh in 4 Cunnel
(diameter of mouth—2?2 cm] leading down
into a glass vial of 75% alcohol, The sumple
was heated from above by a 40 walt electric
light bulb for five days.
Bay samples were collected once a fortnight
for a year (24.1V.1968-23 iv.1969). On each
occasion four samples were collected between
IL am. and 4+ p.m. 2 From the Summit Site
{Sel or S1), and 2 from the Foothills Site
(Fel or Fl)—see appendix, One sumple of
moss on soil and another of plant litter on
soil were taken from each site. A totul of 108
samples weTe collected in the series,
ibe
2 Care Sampling and Extraction
Stee] core samples (5.15 em diameter x 4
cm uepth) were driven into soil covered hy
moss or litter, dug out, and then sealed by a
lid at each end. The volume of euch core was
about 83 ml and from an area of approxi-
mately 20 cm*. The steel cores, without lids,
were inverted On wire mesh in multiple Tull-
gfen funnels, so that the deepest part of the
soil was uppermost, These funncly incorporated
forced draught ventilation to prevent water
condensation, The samples were heated from
above by thermostatically controtled electric
coils to 25° C for 2 days, tollowed by 30° C
for 2 days. 35° C tur 2 days and finally 40° C
for 1 day.
Core samples were only collected on
S.viii.1968 and J2,,ii1.1968, On cach occasion
16 cores were tiken from points evenly spaced
throuvhout cach of 4 plots (S1, $2, FL snd
F2}—see appendix to this paper. Samples from
2 plots (S82 tind F1) were covered hy « suh-
stantial layer of fermenting plant litter, and
the other 2 plots (S1 and F2) were covered
by moss and 4 litle raw leaf litter. A total of
128 core samples were collected in the serus.
Sites
Four siles Werte sampled between the summit
of Mr. Lofty and the coast-line of the Adelaide
Plain, In the appendix, the Summit and Foot-
hills Sites whieh were extensively sampled arc
described in detail, while the Plains and Coastal
Sites Fram which only small collections were
made are given a briefer description, Two plots
at each of the two former sites ure also des-
cribed.
Temperature and rainfall had linear
gradients between the coast and Mt. Lofey: the
former decreasing ond the lattes increasing with
Neurness to the summit. On the other hana, the
Sunimit and Coastal Sites were similar in hav-
ing low nutrient, sandy soils, in contrast to the
higher nutrient, loamy soils of the wo inter-
venient sites. Three sites had a predominately
Jiative Alora, while dhe Foothills Site had an
alien flora,
The sites that were extensively sampled
(Summit and Foothills Sites) included areas
which were elther almost entirety covered by
plant litter or similarly covered by moss, Core
samples were only collected from rectangular
Plots in such Uniforny ures: one moss plot and
one Jitter plot at each of the two sites, On the
other hand bag samples were collected from
larger areas including both plant Iter and moss
patches.
RHODACARIDS FROM NEAR ADELAIDE 14]
RHODACARINAE ¢ : GAMASIPHINAE
143,136 /'60,119
ef Pa
Fae yf \) e.-
te ~ Ww
el aS fh.
eae ve ;
aa i 3 Ly
dk \ f
ee ene i,
Rhodacarus reseus * { \. f
/ oe \ i
ate oad Fa Ne Eveplerius tilamentasus q i
x ;
.
mn
G alistralicus ott
OF (12/00) fs -
«i G 4
f 1 70,33 87,859 ;
v S
¢ i a
Gamasiphis
K a
’ ' 1 :
m ts wa ’ a a mm
r
o 6G fornicatus
¢ 324/ 20.8)
oT
“1
as ; ; G, Jenifornicatus ff
4 te: ¢0.0/5,23 é
Antennolaelapns
OGnchogamasus
virguncula
£0,0/ 7,0)
A
\ 7 ! SESSILUNCINAE
Fig. 1. Species of Rhodacarinae, Gamasiphinae and Sessiluncinae collected during serial sampling,
Dorsal views of adult females, Numbers given under names equal specimens (any stage) from
the following environments: (Foothills Site moss, FS litter/Summit Site litter, SS moss).
142 bD. C. LEE
Geogamasus howardi
0318, 643/41)
OLOGAMASINI
Athiasella relata
695)147/ 52,189
f Axiiassila Jettars “1, OLOGAMASINAE
(503,102 /143,22)
503, 1102/1432 Hiniphis bipala
co,o/s.0)
GAMASELLINI
Solugamasus
mustela
634.7 / 0,0)
Gumaselius
cophinus
¢2,1/97, 9979
&
Gamaselius
tagardh|
(5,0 /321,300)
Ganiasellus concinnis
(5,6 / 451,302)
q
i a \
bo . i Acugamasus puncranis i
Ahodacareides minyaspis Agua BSUS PUNETATS VAcugamasus semIpunctatus’,
(0,0/ 2,3) (0.0/9.0) se 29,0 / 1414)
Fig. 2. Species of Ologamasinae collected during setial sampling, For further explanation sec Fig, [,
RHODACARLDS FROM NHAR ADELAIDE
Resalts
|, Species und /orins represented
Preliminary bag samples from Vhe Plains and
Coustal Sttes in May and June, 1965, produced
the following thodacarids (number of speci-
mens in parenthesis): Plains Site—Rhedacaruy
rosews (18), Gamasiphis australicuy (4),
Athiasella dentata (29); Coastal Site—CGarmua-
sellus grossi (13), Acugemasus elachyaspis
(8)- lt would have been valuable to have pro-
ceeded with scrial sampling at these two sites,
bul becuse of a time limitation this was only
done at the two other sites which supported
more thodacarid species.
Sizes and collection dates of preliminary
samples from the Suramit and Foothills Sites
were not comparable with thase of the other
two siles and are theretore nor Jisted, Serjal
samples from the Summit and Foothills Sites
produced twenty-two species of rhodacarids
(Figs. i, 2), A female Rhadaearetlus silestacas
was listed (Lee 1973) from the Foothills Site,
but this was taken while collecting miles alive
for laboratory cultures. Such collections other-
wise only included species taken in serial
samples.
Only a small proportion of immature chudi-
carids were collected, The 3 species for which
Tesulls are presented (Figs. 6, 7) produced it
relatively high pruporlion of these stages,
Further comments on immature siages are
mide below under Section 5 (Seasonal Varia-
tion),
The sex ratio (male/female) of the 12
commonest rhodacarids From: serial samples is
as Follows; Gamasiphis fornicatus, 0,26: Acu-
gumiasiey . semipunciatux, 0.33, Geegamasus
mininins, 0.344 Geogantasus howarel, 0.41;
Gamasiphis saccus, 0,45; Gamasellus cephians,
0.56: Gamasellus concinius, 0.66; Antes
nolaelaps celex, 0.67; Gamasellus tragardhl,
0.73: Athiasella dentata, 0.83. Euepicrius fila-
meitosus, 0.845 Arhiasella relate, 1.23.
2. Differences between sites
Numbers of specimens were as follows;
Summit Site, 2784 (baz samples, 2)37: core
samples, 647); Foothills Site, 3340 (hag
sumples, 2707; core samples, 633}.
Number ef species were as follows; Summit
Site, 20; Foothills Site, 15. The majurity (13)
of species were foun at both sites, but there
was a considerable difference in the curmpusi-
tion of the fauna. ‘Ihis difference is demon-
straled by presenting the dominance anu fre-
quency of the 12 commonest rhodacarids (Figs.
3, 41. Dominance is the percentage of the total
143
specimens that belong to a species, and is rep-
resented by the size of a shaded urea in a
collimn. Frequency is the percentage of samples
in which a species was found, and is represen-
led by the nuribers in the centre of a shaded
area. Different species are characteristic (i.e.
the most dominant and frequent) of different
sites,
3. Differences between soil cover ef liter
compared with moss
Numbers of specimens were as. follows:
litter, 3558 (bag samples, 2696; core samples,
862). moss, 2566 (bag sumples, 214%; core
sainples, 418). A similar indication is given if
the results from the core samples alone are
expressed as rhudacarids/m? as follows; Sum-
mit Moss Plot, 3450; Summit Litter Plot,
AS60; Fuuthills Moss Plot, 3080; Poothills
Litter Plot, 6310.
Numbers of species from Summit and Foot-
hills Sites were as follows: fitter, 21; moss, 19.
The majority of species (18) were found in
both litter und moss-covered soil. Comparing
the dominance and frequency of individual
species (Figs. 3, 4) it is evident that the species
composition of a particular site is simiar
whether it is cavered by litter or moss. ‘The
preatest differences are shown by the core
samples when the whole plot was mainly
covered by cither litter or moss. Some species
show distinct preferences: e.g. Athiasilla den-
sate for litter; Gamasellus coptinus for moss.
On the other hand. a preference at one site
muy be apparently reversed at the other site,
e.g. Geogamasuys minimis
4. Species Association (only Qlogamusinae)
Exch site has dWferent characteristic species
(see Section 2), mainly belonging to the
Ologamasinae. To eslabhsh whether or not two
communities ate present. the significance of
associations in samples between species of
Ologamasinae is examined. I have followed
Dvebauche (1962) in usin o correlation co-
eflicient based on 4 contingency chi squared
test us a measure of degree of association or
dissociation of species.
The correlation coeflicients fron the results
of bay samples (Table 1) produce the clearest
pattern, Of the 28 terms, 18 ate significunt at
the 1% level or Tess, ‘The species fall into two
groups. One group (afhinselfa dentata and
others) includes species characteristic of the
Foothills Site. The other group (Geagiurasis
tninimus and others) includes species charjc-
teristic of the Summit Site. Since Adhiasella
dentate and A. relata regularly occur al both
ea)
wl
|
U
dD.
144
2)
wi
Oo
=
<
Oo
<
oO
Footwits Summir Summit
Foothills
Species
(Sci+51)
CFci#Fy)
Bhis
Gamasi
is}
o
25959 Rs
OG,
rats
it
fi
Phis
Garnasi
Hews
s
rts oe ky ser fctoeevi re
[damentasus
FuepiéHns
=|
a
entat
a.
howard
Athiaseua
Geogamasus
asus
gam
Geo!
minimus
Gomusellus
sconcinAus
Samasetlus
ophinus
D
Gamaselius
eee
jamasus
Acuy
punctatus
semi
Olaelaps
Antenn
rhodacarids
eclox
Orher
odacarids in bag samples
¥
fs
ge
uo
2
as
im
= 9
2
on
a
w
oo =
Bu
25
=n]
a
eS
we
ar
=
Sa
=m)
=i)
5.
5S
et
Laney
cal
i)
The dominance and frequency of the |
(collected from 2 sites throughout th
145
RHODACARIDS FROM NEAR ADELAIDE
CORE SAMPLES
Summit ‘Summit
Foothills
Foothills
Moss
5
rornicatu
Gamasiphis
seccus
pPicrius
Gus
7
>)
G
°
=
el
DQ
filarm
dentata
Arhiaselia
Athliasella
pS eg Estes ar
fe
aS ce sierra oer STEEISET
eoeeoenee_eetaes ee #ee s® ¢
e_e © 2 68 «Pe eee ee eee Oe
Es
Geogamasus
Geagamasus
minimus
Gamasellys.
SOnclany
Gamasellus
cophinuss:
none
3
Gamesseltu
Acugam.
elaps
Antennola
chnodacsrids
f thodacarids in core samples
the dominance and frequency of the 12 commonest specics o
Fig, 4.
anation see text.
(collected from 4 plots it early August). For further expl
146 D. C. LEE
TABLE 1.
Correlation of Ologamasinae species in 108 bag samples containing 4,844 specimens.
Correlation indices (C X 104), +> (association) .— (dissociation), upper limit is 707.
Species A.den. A.rel. Gow. G.min, G.cop,. G.con. Gira. Asem.
Athiasella dentata +355 +448 —49 —142 —256 —197 —36
Athiasella relata (EY +307 iY) +57 —32 —4$ —t4
Geogamasus howardi Corey ES ee) —319 —495 —583 —S566 —299
Geogumasus minimus 0 0 (4/7) _ +517 +394 = |399 +58
Gamasellus cophinits 0 (///) Cet) +580 +598 +287
Gamasellus cancinnus (f//) 0 (f///) (PFti eer} +656 +396
Gamasellus tragardhi — 0 (//1) (FR) CeRRY (Fea) CR)
Acugamasus semipunctatus ( 0 /f) 0 (ca a Gina +349
Significance of relations:
None Positive Negative
P> 0,1 0
P< 0:1 + =
P< 0.01 (**) (//)
P= 0.001 (ar G//)
the Summit Site as well as the Foothills Site, (Arhiasella dentata—Garnasellus cophinus,
there is, in the main, only a significant dissocia-
tion between Geogamasus howardi und species
al the Summit Site.
The correlation coefficients from the results
of care samples (Table 2) are similar to those
of bag sumples but with 1 drop in significant
associations. Of the 28 terms, 12 (6 associa-
tions, 6 dissociutions) are significant at the 1%
level or less. Although there are the same num-
ber of significant dissociations, 3 are for dif-
ferent pairs of species. The reduction in
associations is only significant where a pair of
species was either uncorrelated or significantly
associated in bag samples while being signifi-
cantly dissociated in core samples. This was
true at the 1% level or less for three pairs
Athiasella dentata—Guamasellus tragardhi, Acu-
gamasus semipunctatus—Gamusellusy Concin-
nus). Such a significant dissociation in core
samples could have had a number of causes:
smaller sample size isolating niches; sampling
separate moss or plant litter covered plots (one
at each site was outside the area used for bag
sampling), thus isolating niches and possibly
introducing new ones: fewer mites per sample;
no seasonal effects such as the absence of any
rhodacarids [rom mest summer samples. The
conspicuous change in dominance between bag
and core sampling of Acigamasus semipune-
ratus In “Foothills moss” samples and Gama-
selius trragardhi m “Summit litter” samples sug-
gests that the significant dissociation in core
TABLE 2.
Correlation of Clogamasinue species in 128 core samples containing 1,280 specimens.
Correlation indices (C X 108), + (association), — (dissociation), upper limit is 707.
Species a.den, A.rel. G.how. G.min, G.cop. G.eon. G-tra. A.semi.
Athiasella dentata +355 +369 —i06 —3&2 —24 —327 —205
Athiayella relata Soaiea | +10 +49 —85 +87 —156 —35
Geogamasus howardi (*#*) 0) =—344 —309 —418 —203 +30
Geogamasus minimus 6 0) (///) +213 +322 +161 —If6é
Gamasellus cophinus (///) 0 f/f) = +264 4236 —102
Guamayellus corcinniy 0 0 (///) (+#*) (er) —20 —231
Gamasellis tragardhi (///) —_ _ + (F*) 0 4355
Acugamasus sermipunctatus _ 0 0 — 0) (//) (**)
Significance of relations:
None Positive Negative
P> 0.1 0
P< 0.1 + —
P< 0.0L (#* (fA
< 0,001 (*??) (//1)
RHODACARIDS FROM NEAR ADELAIDE
samples of the above pairs including these
species is due to the difference in arcas
sampled. The same cause may apply for the
dissociation between Athiasella dentata and
Gamasellus caphinus.
Seasonal Variations
Seasonal fluctuation in total numbers of
rhodaearids at each site based on bag sampling
are summarised in Fig. 5. There was a con-
spicuous fall in numbers of rhodacarids in
samples collected during the summer (Decem-
ber-February). This is associated with a drying
out of the environment and rainfall figures for
Stirling (5 km SSE of Summit Site) are given
as a factor closely associated with this process.
The low number of samples taken means
that seasonal differences indicated for the two
5.
147
sites are tentative. The number of rhodacarids
from the Summit Site was fairly constant
throughout the wetter months (May—Novem-
ber). At the Foothills Site the rhodacarid
population apparently gradually increased in
September and October to a peak, which was
nearly twice the highest number at the Summit
Sitc. If this change is true for the actual popu-
lations it could be telated to the dominant
genus at the Foothills Site, Arkiasella, being
multivoltine, while abundant species at the
Summit Site are univoltine (see below).
Seasonal fluctuation in numbers of the dif-
ferent developmental stages and sexes of
Gamasellus concinnis, Gamasellus tragardhi
and A thiasella dentata are represented by histo-
grams (Figs. 6, 7). Not enough immature
APR MAY JUN Jui AUG sep oct NOV DEC JAN FEB man APA
T 1 | | ] ‘ftw TT T T T
ts a RHODACAAIDS COLLECTED eee
TOTAL pe M--aAT FOOTHILLS SIE —t3a0
|— @--ar SUMMUT SITE hae
NUMBER L. a / ao
an
- . ™ 150
7 /
OF . s a 5
|__ a7 fod
ae \ / on e"-8 a
a Ll s A
fo N x bd
s—a— By a a“
BMECIMENS L. e ‘~,—s 0
‘\ / \
= \ } e PY 150
e ae \
to e—e-* s a \ va \ ran
Jf ‘ ) ‘ a o.
s— 6. at}
— e Na \ 50
X s—e_
i o—*_, ~e fs
RAINTALL AT STIRLING ¢ NEAR SUMMIT SITE)
+ #.—-AcTUAL FORTNIGHTLY fA.
|
| » oy. O---AVERAGE MONTHLY B
Z me
* "Y=
|. y, ee 7,
& [4 eo +
|_ > a
RAINFAL | 4 “sO
“¢ ys,
2 ke af
+ \ ss
IN - \
a , \o
’ 4 bares
“ ls r he re
im 9oNY -~ vo ot)
o L_ + + + \
th ft, =I. | Le al | |
TIME
OF ‘YEAR
Fig, 5. Scasonal fluctuation in numbers. of Rhodacaridae collected in bag samples at two sites during
1968 and 1969. Rainfall records are for nearby Stirling.
148 D.C, LEE
stages of the other rhodacarid species were col-
lected to warrant presentation here.
In Canada, where the winter is extremely
cold, Gameasellus vibrissatus (which is mor-
phologically very similar to G. iragardhi) over-
winters as adult females which give rise to a
single generation in the following summer
(Emberson—footnote 2). My results show that
Gamasellus tragardhi (Fig. 6) bas a similar
life history, except that it over-summers as
adult females and males which give rise to a
single generation in the following winter.
Gamasellus concinnus (Fig. 6) is also univol-
tine, but over-summers in the deutonymph
Stage, the males emerging before the females at
the onset of the wet season, Arhiasella dentata
(Fig, 7) probably over-summers in the adult
stage and it breeds for a longer period, prob-
ubly being multivoltine. Results for some other
species (Lee—footnote 1) are inadequate but
suggest the kind of life-history that they have.
Geogamasus towwardi, Athiaxella relata and
Euepicrius filamentosus appear to have similar
life-histories to Athiasella dentatu. Gamasiphis
vaccus and Gamasellus cophinus may be uni-
voltine and over-summer in the egg or early
immature stages.
It is noteworthy that the life-histories indi-
cate that there are large numbers of rhoda-
carids (e.g. deutonymphs of Gamasellus con-
cinnus) in the soil during the summer that
were not represented in the samples considered
here, Possibly they moye down deeper than the
surface 4 cm sampled. There is no clear indica-
tion that rhedacarid species stagger their life-
histories so as to avoid exploiting the environ-
ment concurrently.
Discussion
The twelve commonest. species of rhoda-
carids in serial samples were found at both
Summit and Foothills Sites, but each site had
ee a y
GAMASELLUS CONCINNUS GAMASELLUS TRAGARDHI
[ FEMALE
i oo
zie
Fr
=
aa
4
a.
a a MALE
no
of
I 7. DEUTON.| is ss
i
ints
ZF
Ee ls ad atrnnd oma Ae tae dans PROTON. eo oh) 79 35 (Eom: oe
=a
=
i
fs
ato eb
APH IMAY! JUIN | JUL) ALS! SER Oct NOY! DEC! Jan] Pea) Man) APR
SE POR VTE ! gba tbat a |
APRIMAY | JUN [JUL | AUG) SEP OCT NOV |DEC. JAN LFEBIMAR APR/
Fig. 6. Seasonal fluctuation in numbers of individuals at different developmental stages of Gamasellus
a
concinnus and G, tragardhi collected in bag samples at two sites during 1968 and 1969.
RHODACARIDS FROM NEAR ADELAIDE 149
= frmale
oe Mate
=
=
2
a
BEUTON,
rm
‘aL,
>
=
3|.
an PROION
mom [tanya
A On sO On
APA MAY ALM dol pale SFP Oa|NoY, Jeo inn Poe lmAR Apr
Fig. 7, Seasanal fluctiation in numbers of in-
dividuals at diferent developmental stages
of Athfasella dentate collected in bag
sanupiss at two sifes duting 1968 and
1969.
different characteristic species and amongst
members of the Ologamasinae there was a
highly significant association between species
characteristic of a site, thus demonstrating the
presence of two rhodacarid communities,
Because serial samples were only taken from
two sites (Summit and Foothills Sites). and
environmental factors were not measured, it is
impossible to confidently associate the rhoda-
carid taxa with particular factors in the envir-
onment. If, however, the few samples from the
Plains and Coastal Sites are considered, there
appears to be a similarity between the rhoda-
carid faunas of the Foothills and Plains Sites,
in that Arhiaselia had the biggest representa-
tlon, and between the Summit and Coustal Sites
in that Gamesellus had the biggest representa-
tion. It appears, therefore, that the coastal sand
dunes as well as the low nutrient, sandy soils
near the summit of Mount Lofty are favour-
able to Garnayellus while the higher nutricnt,
loamy soils of the foothills and plain are not
favourable. The converse appears to be true
for Athiasella. Factors such us temperature and
tainfall, which have a linear gradient between
the coast and the summit of Mount Lofty, do
not appear to be directly favourable or un-
favourable to particular taxa. Tt is noteworthy
that although the flora at the Foothills Site was
introduced, mainly from outside Australia, the
gamasine fauna was predominantly rhodacarid,
with characteristic species that are probably all
eidemic to South Australia and belong ta
genera probably endemic to Australia,
Although the composition of the rhoducatid
fayma of a particular site was similar in soil
samples covered by litter and those covered by
moss. a few species showed a distinct prefer-
ence for one or the other habitat, Other
attributes revealed for certain taxa were the
iendency for species in the same genus to have
similar scx ratios and the specics of one genus,
Gamasellus, to follow quite different life-
histories.
Species of some genera (Arhiasella and
Gamasellus) were characteristic of one site,
while for other genera (Geogarnasus) this was
not true. Conflicting hypotheses on species
association were resolved by Bagenal (1951).
who stated that “related species are more likely
to be found in similar, though not identical,
habitats than are unrelated ones”. Hurlbut
(1968), working on species belonging to
families closely allied to the rbodacarids,
reached a similar conchision expressed as
“species which are very different anatomically
or very similur anatomically coexist less often
than species which are moderately similar to
each other’. Certainly the three species of
Gamaselluy associated at the Summit Site are
as dissimilar (see Fig. 2) from each other as
it is possible to select from known species of
Gamasellus and would probably be considered
by Hurlbutt (1968) as ‘moderately similar’.
They must exploit different ecologtcul niches
within the volume of the small cores in which
they were cqilected. The same is likely.
although not so clear-cut for the two species of
Athiasella,
1
The very slight but easily discernable mor-
phological difference between Gamuavellus tray-
ardhi from the Summit Site and Garmasellus
grossi from the Coastal Site (Lee 1973) sug
gests that the level of taxonomic distinction is
closely associated with ease of anatomical diap-
nosis rather than genetic similarigy. The
Rhodacarus specimen from the Summit Site is
possibly equally dissimilar to the Rioducaruy
specimens From the Plains Site (Lee 1973).
both ef which I have refetred to R. rosens, 1
suspect that many mite species which show
limited distnbution in widely differing geo-
graphical locations (see Introduction) are also
grouped in one species because of difficulties
in diagnosis,
D. C LEE
Acknowledgements
Tam indebted to Dr, K. E, Lee for the use
o£ Tullgren funnels at the Soil Zoology Section
of the C.S.LR.O. Soils Division, and for his
comments on this manuscript, I also acknaw-
ledge the helpful advice afd criticism received
from Dr. D, A, Duckhouse, University of Ade~
laide, during the preparation of the M.Sc, thesis
which formed the basis of this study,
Lam most grateful to Mrs, Curol Aitken for
Preparing some drawings (Figs. 5-7) and to
Mrs: Brenda Head fer preparing the other
drawings (Figs. 1-4) as well as to Mrs, Jo
Bramley and Miss Debbie Rankin for carefully
typing the manuscript.
References
RagewaL, T. Bo 19511—A note on the papers
of Elton and Williams on the generic relu-
lions OF species in smal! ecological com-
munities, f, Anim, Ecol. 20, 242-245.
Dayis, B. N. K. (1963).—A study of micro-
anhrepod communities in mineral soils near
Corby, Northants. J. Anim. Ecol. 32, 49-71,
Denaucnr. Hy R. (1962).---The structural analysis
of Animal Communities of the soil. Jy PW.
Murphy (ed,}, “Progress in Soil Zuvlogy". pp,
16-25. (Butlerworth: London, }
GLYNNE-WILLIAMS, J, & Hopart, J. (1952),—
Sludievs on the crevice fauna of a selected
shore in Anplesey. Prov. 200! See. Lond.
L22, 797-824.
Nurteure A.W. (1968), Coexistence and atia-
tomical similarity In two genera of mites,
Veigala and Asca. Syst, Zoal, 17, 261-27).
| ke, BD, C. (1973)—Rhodacavidae (Acari: Mesas-
tigmats) from near Adelaide, Ausiraliu, 1.
Systematics, Ree. §. Aven Aldus. 16 114), I-
af.
liicHrienp, W, H. (1960).—Soitly on the western
Slopes of the Mt. Lofty Runge near Adelaide
and Elizabeth, South Australia. Divi Rep. Div.
Swils CSIRO 8/59, 1-36,
Luxvon, M, (1967)—The Zonution of Saltmarsh
Acarina, Pedobiolagia 7, 53-6,
Morton, J. E. (1954)—The crevice faunas of the
Upper intertidal zane yt Wembury. J. mar,
bivl, ayy, U.K. 33, (87-224,
Meciak, M., DanteL, M. & Rosicky, B. (1966),—
Parasites and nest inhabitants of small mam-
mals in the western Carpathians, 1. Mites of
the superfamily Gamasoidea (Purasitiformes),
deta Fac, Rerum nat. Waive. eomen.. Brarist.,
Zook 13. 81-11.
SHeaus, J. G. (1957)—The Collembola and
Acaring of uncultivated soil, J. Anin. Eeol.
26, 125-134.
Sencar, R.L.(1972).—"The vegetatinn of South
Australia’. (Government Panter: Adelaide, )
Server, R. £,, & Perey, R, A, (1948).—The
Plant ecology of part of the Mount Lofty
Renaes (1). Troe. Ro Save. 8. Aust, 72, 91-
132.
Woop. T, G. (1967a).—Acari and Collembola of
moorland, soils from Yorkshire, England. |.
Description of the sites and. their populations,
Oikos 1B, 102-117.
Woon, T. G. (1967b).—Acari and Collembola of
moorland soils from Yorkshire, England. 1.
The micro-arthropod communities. Oikey (8,
277-292.
Appendix: Details of Sampling Sites and Plots
Summit Site, Locations Mi. Lofty, near ta sum-
mit, approx. 18 kin trom the sea, Australian Map
Grid vo-grdinales: 290600 nv E/61272320 m N,
map fo. 6628-48-j, Dept. of Lands, Adelaide.
Height above sea level: 640-670m. Rainfall: mean
annual vaintall approx. 120 cm; figures osed in
graph (Fig, 5) are for Stirling (5 km SSE of
site) with a man annual rainfall of 119.0 cm,
and 4 total rainfall in 1968 of 161.6 cm; it should
be noted thal “a large percentage of soinfall is
lost to the soil by run-off an the Adelaide Hills”
(Specht & Perry 1948). Temperature’ mean
monthly min./max, temperatures for Stirling are
January, 11.5°/24.5°C; July, 4,5°/10.5°C. “Ver-
rdiny steep western slope of hill, near to summit,
Soil: Black Hill Assoclation—“low nauicient re-
server im most Soils in Which shallow depth is the
chief limiting physical churacteristic’ (Litchtield
1960); shallow (10-35 em}, dark grey, loamy
sand; an analysis of soil fram Mit, Lofty Summit
showed 0.004469 PO; und 0.0405 Nitrogen
(Specht & Perry 194%). Vegetation: open-forest of
Stringy Burk—Ficwalypiws obliga LoHérit.—with a
sclerophyllous understorey of small native heuth
shrubs including Banksia ornatr FVM. ex
Meisn., EZpucris dipressa Labill. and Leptosper-
mum juniperinum Sam, Fifteen other species of
native shrubs. herhs or grasses were collected from
the site.
Ceneral Snmmreit Plot (Sel). A sub-thombaid
RHODACARIDS FROM NEAR ADELATDP 1S]
area (approximaely 1% a 12 m) which ¢on-
sliluled @ clearing ymanast charred trees with
q draingge channel ronning through the centre
A fire had passed through the plot three years
before (February, 1966). The eastern half of
the clearing had substantial vegetation, includ-
ing abundant fireweed—Sacielia achilleoides
R. Br. ex Ait. a sheart-lived, “high-fertilily-de-
manding” species depending on the temporary
rise in fertiliiy-level due to the ashes of the
burnt vegetation (Specht (972). The western
half of the clearing bad 9 sparse vegetation of
heath shrub seedlings and extensive patches of
moss. Plant Hitter was almost absent from the
mossy half of the clearing, but had wecumu-
lated as raw leaves and twigs around the bases
of tree slumps. Fallen branches and smail shrubs
in the other half, Bag samples were collected
from this plot.
Sammit Moss Plar (Si). A rectangulur area (2
x 10 mj lying approximately at the centre of the
western half of plot Sel, Covered alniat entirely
by on mat of moss. All core samples, and after
August some of the bag samples. were collected
fram this plot,
Summit Liter Plot (S2), A rectaneular area [2
x 10 m) lying approximately 20 m casi of plat
Scl. and separated from It by a bilamen toad,
No fire had been through the plot for 25 years.
Understorey Was thick with heath shrubs und
decomposing plant litter (mainly 1.0 cm deop)
covered most of the ground, Only care samptes
were collected fram this plot,
Foorhills Site, Location: Foothills of My. Lofty.
approx, 16 km from the seu. Australian Map Grid
co-ordinates: 288230 ot E/4127620 m h, map no,
628-49-e, Dept. of Lands, Adelaide, Hetght above
seo level; 240-270 m. Ruinfall: mean annual rain-
fall is appros, 94,0 cm. Temperature; mean
monthly min,/max. temperatures for Glen Osmond
(§ km W of site) ave January, 16°/28°C; July,
7° ¢1S°C, Terrain: hattom of steep northern slupe
(south-Facing aspect of ridge] of Waterfall Gully,
just west of First Waterfall, beside an artificial
pond formed by dredging and damming First
Creek, Soil; Osmond Association—“Nutrient te-
Serves presumably intermediate hetween
law levels ... and the moderate levels in the red
brown carthy af the piedmont aprons” [Litchfield
1940): shallow ta deep (35-110 cm), allavial red-
brown loan, artificially mieved to present position,
possibly from creek hed. Vegetation: alien {is it
mainly i& on bottom 15-20 m of slope downstream
from this point. further up slope from site there
are Manna Giums—Encalypris virinelis var,
huheriana (Naudin) Burbridge-and Drooping
Sheoaks—Caxaerina stricta = Ait}; Fan-leaved
palms (i.fvistonia sp.), Olives (Olea europaea L.),
Lilacs (Syringa wileari« L.) and Fitfesperim (tn
fiefarien Vent, (native of eastern stutes); under-
storce of brambles, bracken und live species of
herhs and grasses; only native plant found was a
small hetb—Gerunium pilosan| Forst,
Gereral Foothills Plor (Fetj. A sub-rectanga-
lar area (approx. 20 © 8 m) which constitited
a patch of quile thick alien vegetalion on the
north bank of the pond, Although the bunk was
steep, the understorey held the plant toec in
nest places. but where the hank was very sleep,
or the ground stony, there was lillle or no
litter and moss or liverworts grew. Bag samples
were collected from this plat.
Foothills Litter Plot (F1). A rectungular urea
(2 © 10m) tying spprex, at the centre of plot
Fel. Covered by plant litter (Olive leaves. pre-
dominated, mainly 2.0 em deep) and some
herbs and eruss tussocks. All core samples, and
after August same of the bag samples, were
collected from this plot.
Foothills Moss Plot (P21, & rectangular qred
(2 x 10 m) lying approx. 10 m west of plot
Fel. und separste! from ic by si artificially
channelled creek bed. Which was stecp and
usually dry. Some soil bare but mostly covered
by moss or liverworts. Under a row of snaall
trees (Pitresporam endalitem) evenly planted
along the west bank ot the creck. Only core
sumples were gollected from this plot.
Plainy Site, Location: Heywood Park, Unley. an
the Adelaide Plain, approx. & km from the sea
Australian Map Grid co-ordinates: 280810 m
E 6128350 m N, map no 6628-50-e, Bem of
Lands, Adelaide, Height above sea level: 30-A0 m-
Rainfall: approx: 38 cm/yeat. Tensperature:
January. [6.4°/29.8°C, July, GS /I45°C, Ter-
rain; small, flat suburban park, wilh fall frees sur
rounding clearing, Sojl; Edwardstoawn Association;
red-brown loum. Vegetation: savannah woodland
of River Red Gums—Encotyptus camatdulensis
Dehnh—"is confined to grey-brown podsols on
jhe slopes and ridges and alluvial soils in the
valleys, both soils being rich in PaO; and nitrogen
and having high water relations” (Specht & Percy
1948): understorey of grass amongst patches of
Fuculyprix liter, Samples fram patches of litter
under River Red Guo.
Coastal Site Location: “Pinery”, Grange Golf
Course, near cost of Adelaide Plain, approx, 1.3
km from the sea, Ausiralian Map Grid co-or-
dimales; 271910 m E/6137D40 m W, map no,
6528-36-m, Dept. of Lands, Adelaide. Height
above sea level: #30 m Rainfall; approx, 43
em/yearm “greens” artificially watered. Tempera-
jure; January, 19°/28°C: July 6.5°/15°C Ter-
rain! inland relicts of coastal dunes formed in
Pleistovene Period, modified to .a golf course. Soil:
Osborne Association; calcareous sand in which the
soluble calcium bicarbonate has been leached to
lower horizons. Vegetation; low woodland of
Native Pines—Callitris preissti Mig.—-with under-
storey of mioss and sparse grase, on dune ridges,
amongst artificial grass wreens; “a “degraded”
climax plunt community characteristic of infertile.
fien-calcareous, sandy snifs” (Specht 1972),
Sumples from mots mats under Native Fines,
st
ase 31 AUGUST, 1973
TRANSACTIONS OF THE
INCORPORATED
CONTENTS
Watson, Jeanette E. Pearson Island Expedition 1969—9. Hydroids - el 5S
Burn, Robert Pearson Island Expedition 1969—10. Opisthobranchs_ - - 201
Seed, W. F. Pearson Island Expedition 1969—11. Crustacea: Isopoda rey,
Daily, B., & Milnes, A. R. Stratigraphy, Structure and Metamorphism of the
Kanmantoo Group (Cambrian) in its YER Section East of
Tunkalilla Beach, South Australia - - - 9G Pails}
PUBLISHED AND SOLD AT THE SOCIETY’S ROOMS
STATE LIBRARY BUILDING
NORTH TERRACE, ADELAIDE, S.A. 5000
—— ee eee we Doe
PEARSON ISLAND EXPEDITION 19697-9. HYDROIDS
BY JEANETTE E.. WATSON*
Summary
WATSON, JEANETTE E. (1973) . -Pearson Island Expedition 1969.-9. Hydroids. Trams. R.
Soc. §. Aust. 97(3), 153-200, 31 August 1973.
Intensive collecting of the sublittoral hydroid fauna of Pearson Island in the Great Australian Bight,
in January, 1969, yielded 81 species (with 3 varieties in one species), of which 13 species are newly
described. There are 18 new records for South Australia and 1 new record for Australia. The
collection permits a fuller description of seyeral hitherto poorly known southern Australian species.
Collections were made using SCUBA at 3 localities representative of environmental extremes on
the coastline-a rough-water site exposed to prevailing swell, a sheltered embayment, and a deep
water situation in open ocean.
The deeper water fauna contained species already known from deep dredgings in the
Great Australian Bight, but differed markedly from the collection from shallower water, with only 1
species common to both.
The Sertulariidae and Plumulariidae are represented by the greatest number of species, and are
equally abundant in both epizoic and epiphytic habitats; the Haleciidae, Lafoeidae and Syntheciidae
are epizoic, and Lineolariidae, with 1 species, epiphytic. The large plumose colonies of the
Aglaopheniinae are epilithic. Hydroids are more abundant on the rough-water coastline, where red
algae and the solitary ascidian, Herdmania momus (Savigny), are epiphytised by a large number of
species. Delicate athecate species, and species of the Campanulariidae which may be expected to
liberate medusae, are restricted to sheltered waters, or to depths below turbulence from surge in the
rough-water locality.
The high percentage of hydroids now known to be common to the coasts of South Australia,
Tasmania and Victoria, supports the view that the Flindersian proyince extends from Bass Strait
into the Great Australian Bight,
PEARSON ISLAND EXPEDITION 1969+—9. HYDROIDS
by JEANETTE E. WATSON*
Summary =~
Warson, Jxanetre E. (1973)—Pearson Island Expedition 1969,—9, Hydroids. Trans. R,
Sec, 8. Aust. 97(3)_ 153-200, 31 August 1973.
Intensive collecting of the sublittoral hydroid fauna of Pearson Island in the Great
Australian Bight, in January, 1969, yielded #1 species (with 3 varietics in one species), of
which 13 species are newly described, There are 18 mew records for South Australia and 1
new record for Australia. The collection permits a fuller description of several hitherto poorly
known southern Australian species, :
Collections were made using SCUBA at 3 localities representative of environmental
extremes on the coastline—a rough-water site expased to prevailing swell, a sheltered embay-
ment, and a deep wuler situation in open ocean.
The deeper water fauna contained species already known from deep dredgings in the
Great Australian Bight, but differed markediy from the collection from shallower water. with
only 1 species common to both,
The Sertulariidae and Plumoulariidae are represented by the greatest number of species,
and are cqually abundant in both epizoic and epiphytic habitats; the Haleciidae, Lafoeidae and
Syntheciidae are epizoic, and Lineolariidae, with 1 species, epiphytic. The large plumose
colonies of the Aglaopheniinae are epilithic. Hydroids ar¢ more abundant on the rough-water
coustline, where ted algae and the solitary ascidian, Herdmania maomus (Savigny). are
epiphytised by a large number of species. Delicate athecate species, und species of the Cam-
panulariidac which may be expected lo liberate medusae, are restncted to sheltered waters. or
to depths below turbulence from surge in the rough-water Jocality.
The high percentage of hydroids now known to be common to the coasts of South
Australia, ‘Tasmania and Victoria, supports ihe view that the Plindersian preyince extends
from Bass Strait info the Great Australian Bight,
Introduction
Hydroids have been reported from a num-
ber of expeditions around the southern and
south-eastern Australian cOastlineé—the yoyage
of the “Rattlesnake” (Busk 1852), the “Chal-
lenger” dredgings in Bass Strait (Allman 1883,
(884), the “Thetis” dredgings along the New
South Wales coastline (Ritchie 1911). the
“Findeyyour” expeditions from New South
Wales to Western Australia (Bale 1914, 1915),
the Michaelson-Hattmeyer Expedition to
Western Australia (Stechow 1924, 1925}, and
the McCoy Society Expeditions to Lady Julia
Percy Island, Victoria (Blackburn 1937), and
the Sir Joseph Banks Group, South Australia
(Blackburn 1938). With the exception of the
last two expeditions, the hydroid collections
were made over a wide geographical area,
while the collections of the Michaelson-Hart-
meyet Expedition and the McCay Society Ex-
peditions, although restricted in urea, were
gained mainly fram drifi, the eulittoral zone,
and to a minor extent, from shallow subtidal
dredgings. No intensive survey has. however,
been made of the subtidal hydroid fauna at
any one locality in the Australian region,
The joint expedition of the Department of
Fisheries and Fauna Conservation of South
Australia and the Royal Society of South Aus-
tralia to Pearson I, 6-15 January, 1969, pro-
vided an opportunity to undertake a compre-
hensive subtidal survey of the hydroid fauna of
an offshore island.
“Honorary Associate, Invertebrate Zoology, National Museum of Victoria, Melbourne, Vic. 3000.
+ Other accounts of the geomorphology and biology of the Pearson Islands are given in Volume 95,
Part 3 (4971) of the Transactions, as well as in the present part.
1%4
The South Australian hydroid fauna is
known from reports of the ‘‘Endeavour™
dredgings in the Great Australian Bight along
the 126° parallel of longitude, together with a
small collection from the Isles of St. Francis
in the Nuyts Archipelago, Blackburn (1938)
recorded the shallow water fauna of the Sir
Joseph Banks Group in Spencer Gulf, and
Shepherd & Watson (1970) listed and dis-
cussed the associations of hydroids and algac
at West 1., Encounter Bay.
The survey yielded a total of 81 species
(with 3 varieties in one species), of which 13
species ate new; there are 18 new records for
South Australia, including { new record for
Australian waters. Only 2 athecale species
were found, and 6 of the thecate hydroids
could be identified only to genus.
Pearson I. (Fig, 1) is:a graniuc island situ-
ated ai Lat. 33°57'S, Long. 134°15'E. about
shelfered site
roar
he
Eastern Cove
Lang J34° 15'E
Lot 33° 5a! go"s
NORTH SECTION
Harte
wf aval
SECTION
2 409 8
metres
Fig. 1. Map of Pearson Island, showing study
sites near the northern cnd. Inset shows
the situation of Pearson Island in the
Great Australian Bight. (After Shepherd
& Womersley 1972).
IRANETIE &, WALSON
64 kim offshore on the continental shelf in the
eustern fegion of the Greut Australian Bight,
Weatheritig of the granite has produced a
rugged Lopography of massive blocks, cletts
aud caverns, continnous to the seafloor at
depths of 45 m immediately surrounding the
island, with a rapid increase in depth offshore
to 70 m. A more detiled account of the en-
vironmental conditions is given by Shepherd
& Womersley (1971),
Methods
Collections were mude by divers using
SCUBA. As diving time was limited toa total
of 30 hours underwater, two main sites were
chosen for intensive collecting. One site wus
on the rough-water windward, southwesterly
side of the island; the other was in the more
sheltered north facing Eastern Cove (Fig. 1),
The benthic flora and fauna at each sile was
systematically sampled (with particular atten-
tion to hydroids) from the upper sublittoral
10 the seafloor at 50 m depth. Two additional
small epllections were also made—one in sea-
grass meadows in the more sheltered part of
Exsiern Cove, und another, at 65 m depth, 4
km to the south, between Pearson 7. and
Dorothee (Station F), Because of the rugged
nature anc exposure of the coastline to surf,
no collection of the intertidal fauna was made,
Collections
Holotype and paratype microslides, and
other microstides and material are lodged jn
the Nutional Museum of Victoria, Melbourne
(NMY), Paratype microslides are also lodged
in the South Australian Museum (SAM).
In most instances, the synonymy of Rulph
(1958, 196la.b, 1966) is adopted, and only
pertinent references to species in Australian
literature ure given. The status of several
species is reviewed to tesalye confusion in the
literature, and a number of rare and poorly
known species ate redescribed,
The site notation of Shepherd & Womersley
(1971) is followed, “R” denoting material col-
lected on the rough-water side of the island,
and “S” denoting the sheltcred side. Hydroids
collected in deep water at Stalion F are noted
separatcly. Depths at which cach species was
collected ure given. These depths will, how-
ever, represent only part of the total range of
each species. In most instances, the substrate
upon which each species was found is ulso
noted, As many hydroids ate hoth seasonal
and irregular in occurrence, it is likely that
collectiuns made at oiher parts of the island
HYDROIDS
or even at the same sites at other times, would
yield a slightly different faunal list.
The algal acology of these Pearson L. sites
is described by Shepherd & Womersley (L971),
who also list the algal species collected.
Ecology
Oceurrence af Hydroids
The collection yielded a large number of
species, in spite of the apparent paucity of the
hydroid fauna on Arst inspection of the locality.
Many of the species are crypuc forms growing
sparscly In small colonies among algae, where
only accessible with SCUBA. Even those
species (Selanderia fusca, Thecocarpus divari-
cates var. maccoyi, Aalicorearia longiroxtris,
H. prafifera) which are known to form con-
spicuous colonres in other localities, Were small
and attenuated, suggesting that conditions for
growth were not entirely favourable. This may
be due to the strong surge conditions around
the island preventing good grawth of the
colonies, or to the high light transmittance of
these waters, which appears to inhibit bydroid
growth (J. W. unpublished data).
The only athecate species, Tuhuleria larynx
and Selanderia fuseu, recorded from Pearson
I, were Found in relatively sheltered situations
on the rough-water site. Only one colony of
the former species, growing deep within a
euvern, was recorded, whereas the Jarter
spevics occurred as abundant small colonies
in sheltered situations among the holdfasts of
the brown kelp Ecklonia radiara.
The scarcity in rough ocean waters. of athe-
cate and other specics which liberate medusae
has ulready been noted at West, I. (Shepherd
& Watson 1970). Thua their absence from
Pearson I. is not surprising, considering the
exposure of the coastline to surf. Caripanu-
faria wustealis, a species likely to liberate
medusae, but whose reproductive strictures
ure still unknown, was however common on
the rough-water site, but only at a depth of
34-30 m, well below the zone of maximum
turbulence,
The collection from Station F, at 635 m
depth, yielded a markedly different fauna from
that of shallower water. The seafloor at this
station was tippleanarked sand, with tare
algae, a sparse epifauna of worm tubes, cal-
caréous hryozou, solitary acidians, and ald
shell. The dominant hydroids here were Syen-
plectoscyphus subdichotomus, S. fongithecus,
Syrtheciuin elegans £. subventricosumt and
Plamularia axsymmetrica The only species of
this group also fanging into shallower water
15
was S. slbdichoionius bit it Was uncommon,
P. asyrumetricg, the most abundant species in
the deeper water, has heen recorded only
three times previously, from adjacent waters of
the Great Australian Bight.
Reiaionship berween Hydroid anid Subsinate
Although little firm evidence of the associa-
fon between hydroids and substrate can be
gaincd on the hasis of ane series of collections,
a mumber of broad relationships aud possible
obligatory associutions are evident from the
Peurson [. material.
OF the total of &1 specres and 3 varieties, 27
urc exclusively epiphytic, 30 are non-cpiphytic,
und 18 are both epiphytic and epizoic, The
holdfast fauna, although strictly epiphytic, is
listed with epizoic, epilithic and "{ceegrowing”
species. (Table 3.)
A. Bpiphytie Hydroids (Table 2).
The two major families present, the Ser-
tulariidae and the Plumulariidue, are almost
equally divided between epiphytic and ¢pizoic
species. The Lineolariidae,. represented only by
Lineolaria spinulese, is epiphytic, Altogether,
47 species ate associated with red algac. mostly
on the rough-water sile, 27 species are associy-
ted with brown algue, and 3 species with green
algue, This is tn accord with findings at West
f. (Shepherd & Watson 1970) where red algae
were alsa the most heavily epiphytised group.
The most abuadant and widely distributed
species of algae had the greatest number of
hydroid epiphytes. Brown algae, particularly
Sargassum, although a substrate for fewer
hydroids, were often heavily epiphytsed by
luxuriant colonies of some common apecies
such as Amphishelia mininia var. pumiloides,
Plumularia epibracteolosa and = Sertularella
avrilia.
Several hydroids showed a hich degree of
selectivity and were ussociated with only ane
algal substrate (Table 2}, but as these species
Were not of common occurrence, it i uncer
tain whether an obligatory relationship actually
exists. However, there are two inslances of a
commonly occurring hydroid associated with
only one species of alga. Sertwaria acuta was
found only on the red alga Stenoeliadia aus-
tralix, whereas at West [. it wis abundantly
associated with the red alga Piacelecarpus
lahilfardier’. ‘The factors influencing the pref-
erence of §. acura for Stenocludia atustralfy, to
the exclusion of PRacerecarpus labitlardieri, at
Pearson I. are unknown.
The mast vonspicgtious assaciatinin Was thut
of Plamualarte epibructeoiosa with Use brown
ISG JEANETTE bE WALSON
alea Sargassum bracreolosuin, The fronds of
this alva were densely covered hy the hydroid,
in contrast to the stems which were exclusively
epiphytised by Amphishetia snininta vat.
puniloides. S. bracteolosunr was also recorded
at West |., but was epiphytised only by A,
aint and Campanularia ausivaliv.
Two other species, Lineolaria spinulose and
Plorindaria dastralis, were common epiphytes
tm the séagriss Posidonia australis, but were
not associated with algae. Colonies of both
hydroids were Frequently intergrown on, the
same blade of seagrass.
The growth habit of Phunuluria flexuosa
with a species ol the ved algal genus M yehedea
has not previously been reported, Usually
hydroids avoid algae with aw thallus of small
diameter (Nishihira 1967), The frond of
Muochodee sp., although somewhat Jurger than
the hydroid stolon, js nevertheless rather nar-
row. Vhe hydroid stolon passes length wise in-
letnally through the frond, giving off stems
to the outer surface at regular intervals.
Of particular interest is the gradational
epiphylism displayed by the Arniphisbetia
mininia-A, minuseule group, Bale (1884) dis-
tinguished var. pumtiloides from, var. intermedia
entirely upon the structures of the trophosome.
Hoth these varieties, and the closely related
species A. minusent, are abundant in the
present collection, und display a marked
gradational preference for certain groups of
ulgae—the robust var, pumilaides is found on
larec brown algac of the genus Sargassum (8.
varjans, §. verruculosion, S. bracteolosumy and
On Acrecerpia paniculaa; var. intermedia, a
small Lorm, is assuciated with the red algae
Rhodvinenia australis, Metamaytophera fabel-
lata, Levrencia élata, and Carpopeltis pipvllo-
Phord A, minuscula is both epiphytic and epi-
zuic and is associated with the red alga Lavrea-
cia elare, the brown alga Distromtium flabetle-
tum, a species of the green alga Caulerpa, as
well us being epizoie on Herdmania monnes
and Aalicornaria longirosiris,
B. Epizoic Hydroids
The Lafocidac, Syntheciidae and Halectidae
(with one exception Halecium sp. 1) are en-
tirely eépizoic. Generally, epizoic associations
are less well defined than epiphytic assacia-
tions, most of the species involved being found
on a wide variety of animal substrates, The
substrates upon which hydroids: were found
were, in order of abundance, other hydroids
(19 oceurpences), culcureous bryozou (15),
splines (4), the sulitary ascidian Herdnanig
mommies (l4), compound aseidians (3). und
muciliginous worm tubes. (2). Of the animal
Substrates, only A. moms and other hydroids
could be identified to species,
The two species of hydroids most commanly
enpizoitised by other hydroids are Thececarpas
divaricatus var. eystifera and Halicormmaria
longirastris, whose thick robust stems are sui
able for colonisation by the small stolonic.
species such as Syvaiplecroscyphus epizoicns,
Retieularia antarctica and R. gsnulare,
Herdmania rromus, one of the most ahun-
dint larger invertebrates at Pearson I,, grows
upon rock walls in open situations where there
is modetale water movement. The leathery
siphonal region of the ascidian is colonised by
small species of red algae, bryozoans and hy-
droids, the most commonly occurring hydroics
being Sertulerelle robusta, Sertularella sp. 1,
and Diphasia siebcar late.
C. Epilithic Uydroids
Epilithic colonies are usually conspicuous
plumose forms growing from small matted
routstovks wn rock surfaces in open situutions
where they can take: maximum advantage of
Water movement, This group, comprising
Thecocarpuy divaricaus val, eystifera, Malicey-
naria profifera, H. fonpiroyrris, and H. aurea,
all belong ta the Aglsopheniinae,
Closely allied in habit ta the true cpilithic
species are the lwo “freegrowing™ plumularian
species, Plumularia asymmetrica and Halop-
teris xtleda. Both wre large plumose colonies
growing from a small rootstock attached to
pebbles or shell fragments buried in the seu-
foor,
D. Ner-selective Hydroids
Species oceurring in both epizvic and epiphy-
lic habitats are often also associated with the
most ubunodyot animals ind plants, and are
thus amoung the most commonly occurring
hydroids, The most frequent of these multi-
preferential associations arc:
Hydeaied
Sertularella rohusra
Srebstrate
Ballia callipricha
Herdmania momus
Cavreneta eluta
Herdmania momus
Amenysla pinneri fides
Herdmania morass
Sertutwrelia sp 1,
Diphasia subcarinate
ee ee ee
Although many species (18) are both epi-
zoic and epiphytic, only 2 species, Thecocarpus
divericunsy and Halicarnaria longirostris are
epilithic as well. The largest of the 3 varieties
of FT. divuricetus, var. crstifera, is epilithic.
HYDROIDS 157
while of the 2 smallee varieties, var, 5rigesi is
pradational between epiphyde and epizoic, and
var, macceyi is epiphytic.
Distribution
All thecate families, with the exception of
the Campiunuliniidae, are represented in the
collections. Campanulariidae are represented
with 5 species, Sertulariidae with 31, Plumu-
lariidae with 31, Halecitdae with 5, Lafoeidae
with 4, Syntheciidie with 2, and Lincolanidac
with | species
With the exception of Thecocerpus divari-
cutus var. drives’ (previously recorded trom
New South Wales), ARericwaria antaretica
(Western Australia), and Zygophylax anti-
puhes (Torres: Strait), all other species newly
recorded from South Australia are known trom
Victorian waters, mostly from the intensive col-
lecting of Mulder & Trebileack (1909-1916)
along the central Victorian coustline, from Port
Phillip Heads to Torquay,
The new record for Australia, Syatheeiusa
dentigerum, has been reported only twice pre-
viously, once from the Indian Ocean, und once
from South Africa.
The genus Lyrocarpus, well known from the
Indo-Pacific region, ts recorded for the first
time (1, nuléer’) trom southern waters.
Only 14 of the 38 species listed by Black-
burn (1938) from the Sir Joseph Banks Ciroup
were in the Pearson I, collections. The two
groups of islands, however, are subject to dil-
ferent environmental conditions, the former
group of islands being situated in sheltered
water at the southern end of Spencer Gulf,
in contrast te the extreme exposure to rough
water of Pearson I. in the Great Australian
Bight. Comparison of the faunal lists of the
twa island groups shows thal the species com-
mon to both ure mostly specics epiphytic on
algae, and on the seagrass Pasidenia casusiralts.
Zoogeogruphy
Of the 83 specics and 3 varictics recorded
from Pearson |. 18 spectés are comnion to
New Zealand waters, 15 have a northern Aus-
tralian and western Indo-Pacific distribution,
10 specics occur jn South African waters, 6
are recorded from Japan, 2 from the Antarctic,
and 2 are cosmopolitan. Thirty-nine (49%) of
the species recorded (including the new
species) ate, us presently known, cndemic to
southern and south-eastern waters of Australia.
Blackburn (1942) estimated that 42%. of
the known hydroid Iouny of South Australia
Tanged into New South Wales, and 189% into
Western Australia, The present collection (with
1 variety common to N,S,W. and 2 species
common to W. Aust.) does not substantially
alter these estimates, The basis of comparison
between the 3 States is. however, poor, as both
the Jeep and shallow water hydroid fauna ot
South Australia as now better known from
SCUBA collections, whereas much of our
knowledge of the hydroid fauna of New South
Wales comes from deeper dredgings On the
continental shelf, and that of Western Australia
is from the reports of shallow water collections
between Albany and Shark Bay (Stechow
1924, 1925).
The 18 new records from the present colfec-
tion, combined with 18 from West T, (Shep-
herd & Watson 1970) brings the South Aus-
tralian bydroid fauna, based on Blackburn's
list, to 1 L9 species.
Thus, the total number of species common
to South Australia and Victoria is 81 (61%)
of the knewn South Australian fauna, This
fizure docs not differ greatly from Blackburn's
earlier estimate of 65%,
Based on Hodgson (1950), the fuuna com-
mon to both South Australia and Tasmania ts
69% of the South Australian fauna. Che
species common to Victoria and Tasmania
comprises 70% of the known Tasmanian
fauna. This distribution putlern lends further
support to the contention of Womersley & Ed-
monds (1958) that the Flindersian Province
embraces much of the Maugean, anil exterils
from the eastern Vicrorian coastline io at Irast
the central coastline of South Austritia.
Systematic Section
Order ATHECATA
Family TUBULARTIDAE
Tobutaria larynx Ellis & Solander, 1786, 31,
Bule, 1888: 748. Ralph, 1953: 68; 1966;
160.
Reeords: R, 24 m, on walls of cavern, shel-
tered from surge.
Material: One small cluster of stems to 2 em
high, Sven increasing gradually im diam, dis-
tally ts OS em. Perisare thick, smooth, wilh
groups of 3-8 annulalions; regrowth of broken
stems bepinning with a new series of annula-
tions. Hydranth 1.2 mm long, 1.0 mr wide,
but tentacles not fully extended. Proximal
whorl of tentacles a Jitthe longer tha clistul.
Gonophores small, spherical, sex tideterminate,
clustered between whorls of tentucies. Celoii—
tentocles white, gonophores pink
158 JEANETTE E. WATSON
TABLE 1
List of Species
Substrate notation; EZ = epizoic, Ep = epiphytic, El = epilithic, Fg = freegrowing, Hf = holdfast
fauna.
Symbols are given In order of abundance of colonies on substrate,
* denotes a new record for South Australia. .
The number preceding names of the species in the following list is the key to the species in Tables 2 and
ATHECATA
Family TUBULARITIDAE
1. *Tubylaria larynx Ellis & Solander, El.
Family SOLANDERIIDAE
2. Solanderia fusca (Gray). Hf,
THECATA
Family CAMPANULARIIDAE
3. Clytia (?) pearsonensis n.sp. Ez.
4. Campanularia ambiplica Mulder & Trebileock. Ep.
5. Campanularia australis Stechow. Ep, Ez.
6. *Campanuluria gaussica Stechow. Fz,
7. Campanularia sp. Ep.
Family LAFOEIDAE
8. *Reticuluria entarctica (Hartlaub), Ez.
9. Reticularia annulata n.sp. Ez.
10. Reticularia sp, Ez.
11. *Zygophylax anfipathes (Lamarck). Hf.
Family LINEOLARTIDAE
12. Linevlaria spinulosa Hincks. Ep.
Family WALECUDAE
13, *Ophiodissa australis (Bale). Ep.
14, Ophoidissa blackburni nsp. Ez.
1S. Phylactotheca armata Stechow. Ez.
16. Halecium delicalulum Coughtrey. Ez.
17. Halecium sp. 1. Ez.
18. Halecium sp. 2. Ep.
Family SYNTHECIIDAR
19, Synthecium elegans forma subventricosum Bale. Ez.
20. “Synthecium dentigerum Jarvis, Eu
Family SERTULARIIDAE
21. Thyroscyphus marginatus (Bale). Ep, Ez.
22. Parascyphus simplex (Lamouroux), Ez.
23, Diphasia subcarinata (Rusk). Ep, Ez.
24. Stereotheca elongata (Lamouroux), Ep.
25. Crateritheca acanthostoma (Bale), Ep, Ez.
26, *Crateritheca crenata (Bale). Ep.
27. Sulucia obliquanoda (Mulder & Trebilcock), Ep, Ez.
28. Sertularella robusta Coughtrey, Ep, Ez.
29. *Sertilarella simplex (Hutton). Ez.
30. Sertularella annulaventricosa Mulder & Trebileock Ep, Ez.
31. Sertulareila avrilia n.sp- Ep.
32. Sertularella sp. 1. Ez, Ep,
33. Sertularella sp. 2. ?
34. *Symplectoxcyphus longithecus (Bale). ?
35. Symplectoscyphus subdichotomus (Kirchenpauer). Ez.
36. Symplectoscyphus neglectus (Thompson), Ep.
37, Symplectoscyphus indivisus (Bale). Ep.
38, Symplectoscyphus pygmaeus ?(Bale), Ez.
39. Symplectuscyphus macrothecus (Bale), Ep.
40. Symplectoscyphus rostratus n.sp. Ep, Ez.
41. Symplectoscyphus epizoicus n.sp. Ez.
42. Sertularia macrocarpa Bale. Hf.
43, Sertularia iinguicilata Busk, Ez, Hf.
44, *Sertularia bicuspidata Lamarck. Ep.
45, Sertularia maccallumi Bartlett, Ep.
46, Sertularia acuta Stechow, Ep.
47, Amphishetia maplestonei (Bale). Hf.
48. Amphisbetia pulchella (Thompson). Ep, Ez.
49, Amphisbetia elyeni n.sp, Fz, Ep,
HYDROINS 159
50. Amplishetia minima var. pumiloides Bale, Ep,
St. Amphixhetia minima var. intermedia Bale. Ep. Ez.
32. Amphishetia minuscula (Bale). Ep, Ez.
Family PLUMULARTITDAK
33. Pyenotheea producta (Bale). Ep.
54. *Amtennella tubulosa (Bale). Ep
“Antennella campuniliformis (Mulder & Trebilcock). Ep.
56. *Antennella secundaria s.sp, dubiafermis (Mulder & Trebilcack ), Ep,
37. Halopteris suleata (Lamarck). Fe.
38, Halopteris campanila var. campanula (Busk). El,
59. Halapteris biuski (Bale), Fz.
60. Halopteriy oppasita (Mulder & Trebileack). Ep.
61. *Gaitva bale (Bartlett). Ep.
62. Gattya uglaopheniaformis (Mulder & Trebilcock). Ez, Ep.
63. Gaftya trebileovki nusp. Ep.
64. Plumularia procumbens Spencer. Fl.
65. Plumularia asymmetrica Bale, Fg.
66. Plumutaria flecnosa Bale. Ep.
67. Plumularia spinnlosa Bale, Ep. Ez.
68. *Plumularia goldsteini Bale. Ep.
69. Plumularia obliqua (Johnston). Ep.
70, Plumularia australis Kirchenpaver. Ep.
71, Plumutaria epibractealosa n.sp, Ep.
72. Plumularia meretricia asp. Ez.
73. Plurmitlaria togata n.sp. Ep.
74, Plimularia australiensis n.sp. Ez,
75, Aglaophenia plumosa Bale. Fz, Ep.
76. Thecocarpus divaricatiis yar. maccayi Bale, Fp.
77. *Thecocarpus divaricatus var. brigest Bale. Ep, Ez.
78. Theeocarpus divaricutys var. cystifera Bale. Fl.
79. *Lyvtocarpus mulderi (Rartlett). ?
80. Malicarnopsis elegans (Lamarck). E}.
81. Halicornaria longirastris (Kirchenpauer). FJ, Ez, Ep.
82. *Halicornaria prolifera Bale. El.
83. Halivernaria aurea np. EL
Remarks; This cosmopolitan species was douht-
fully recorded for the first time from Austra-
lian waters by Ralph (1966) who reported a
few infertile stems from Port Phillip Bay, Vic,
This is the second tecord of the species in Aus-
tralia, and a new record for S. Aust.
Solanderia fuseq (Gray, 1868). Watson é& Uti-
nomi, 1971; 19, pl. 8.
Ceratella fusca (Gray), Spencer, 18915 &-
Records: R, 14-33 m, among holdfasts of
brown algac.
Material: Four very small infertile colonies
broken off from the rootstock, the largest
colony 5S mm high and 20 mm _ wide.
Colonics compact. branching closely in one
plane from a thick main stem, Srem of largest
colony 3 mm wide at base, stem and branches
flattened in plane of growth. Aydrophores are
Open shelf-like structures, prominent on
younger branches, edged with 10-15: bluntly
pointed terminal spines connected by a thick.
shallowly scalloped chitinous web. Trabecilate
meshwork of branches close and solid, with
square to circular openings, Spires similar to
those edging hydrophore developed at points of
intersection of meshwork on older branches.
iTvdranuths poorly preserved. Colour—stems
dark brown, shading to ight brown on grow-
ing tips.
Remarks: The colonies of §. fitsce from Pear-
son J., although dwarfed and infertile, are
mature, occurring among algae on horizontal
rack faces. This. is in contrast to the known
habitat of larger specimens from Victorian
waters, which seem to favour vertical walls
and the interior of caverns (J.W., unpub-
lished). Bale (1888, p. 749) mentions that his
small colonies from Sydney were from “Lamin-
aria roots” (probably Ecklonia radiata hold-
fasts). Watson & Utinomi (1971). reported that
the spinous trabeculae were not present in
material examined by them from the Great
Australian Bight, vet the Pearson J. specimens
show these spines clearly. Unknown environ-
mental and geographical factors may thus in-
fluence structural variations within the species.
Order THECATA
Family CAMPANULARIIDAE
Clytia (’) pearsonensis n.sp_
FIG. 2
160
JEANETTE E. WATSON
TABLE 2
Fpiphyic Hydroiils
The numbers tefer to the species as given in the species list.
VC = very common, C = common, R = rare,
Aleal Substrate Hydroid
CHLOROPHYTA
Canlerpa brownii (C.Ag.) Endlicher. 63(R)
Cailerpa staplicinsenle (Turner) J. Agardh. T7(R)
Caulerpa sp. 52(VC)
PHAEQPHYTA
Disrominm flabellaium Womersley. §2(VC), 53(R), 56(R)
Distramium sp. 37(C)
Zonaria spiralis J. Agardh. Ta(VOY
Sevtotlalia darvcarpea (Turn.) Greville. 24(VC)
Acrocarpia paniculala (Turn) Areschoug. 24(VC), 39(R), S0(VC), 76(VC)
Cystephora brownii (Turn, ) J, Agardh, 5(C)
Sargassum verruculosum (Mert-) I Agardh. 31(C), 40(C). 60(R)
Sargassum hractéolesum J. Agardh. S0(VC),7I(VC)
Sargassum spinuligerum Sonder,
Sargassum Varians Sonder.
Sargassum 8D,
RHODOPHYTA |
Delisea pulehra (Grev.) Montagne.
Preracladia lucida (R.Br.) J. Agardh.
Rhodopeltis anstealis Harvey.
Metazoniolithon chargides (Lamx.) W. v. Bosse.
Metamasiophara flabellata (Sond.) Setchell,
Carpopeltis phyllophara (A. & A.) Schmitz.
Callopha lis caccinea Harvey.
Plocamium aneastune (J, Agardh) Hooker & Harvey.
Hacamium cartilagineum (L.) Dixon.
Phacelocarpus labillardieri (Mert.) J. Agardh,
Stvenoclacdia australis (Sond.) Silva.
Alvehodea sp,
Rhoadymenia anusiratis Sonder.
Ballia callitricha (C.Ag,) Kuetzing.
Prerosiphonia?
Anunsia pinnatifida Harvey.
Laurencia clata (C.Ag.) Harvey.
Inidentified red algac.
ANGIOSPERMAE
Pasidonia australis Hook. f.
37(C), 50(VC), 60(R)
sa.VC), 55(R)
21(R), 30(C), 36(VC), I7(C), S4ER), 69(R)
48(R)
5(C), 36(VC), 45(VC). 48(R), 55(CY
61(C)
44(R)
27(VC), 36(VC), 73°VC)
44(R), SICVC), 61(C), 69(R), T6(VCO)
45(VC). 75(R)
62(R)
67(C)
7T6(VC)
24(VC)
24(VC), 46(VC)
66(R)
SI(VC), 56(C)
28(C)
61(C)
23(VC)
32(VC), Si¢VC), S52°VC), SS(C),
67(C)
4(R), 70K), 25(R). 27 (VE), 30(€). 36(VC).
40(R), 49(C), 53(R), T7(R), STC VO)
12(VC), TOCVE)
37(C),
Type Material and Records; Holotype,
NMYV G1914, microslide—R, 22 m, on stem
of Thececarpus divaricatis var. cystifere;
paratype, G1915, microslide—R, 34 m, on
bryozoa.
Description from holotype and paratype: Pedi-
cels long, of variable diametcr. irregularly
wrinkled or smooth (holotype shows indistiact
distal annulations), arising from a creeping
stolon. Hydratheede large, cylindrical. walls
smooth, perisarc very thin and delicate, mar-
vin entire. everted into a thin lip. Thecal wall
slightly thickened proximally, hydrotheca with-
out floor, tapering inta pedicel, a trace of a
very thin diaphragm near base. Hvdranth too
poorly preserved for diagnosis, Ganotheca ab-
sent,
Remarks: Only 2 undamaged hydrothecae were
found im the entire collection, although othet's,
badly damaged, were noted. It is possible that
the species has previously been overlooked
hecause of the extremely delicate nature of the
perisare, which collapses immediately on re-
moval from water.
C. pearsonensiy ts closely related ta Lao-
medea michael-sarsi Léloup, 1935, reported
from only two localities—the West Indies, and
the west coast of North Africa. The hydro-
theca of L. michael-sarsi are, however,
shorter and less than half the diameter of C.
pearsonensis. (Measurements ate given for
camparison),
Following Millard (1959, p. 248), the
species may be referable to cither Campatu-
HYDROIDS 161
TABLE 3
Epizoic, Epilithic, Freegrowing and Holdfast Associations.
Substrate
Wydroid
Epizoic Species
Ascidian— .
Herdmania momus (Savigny)
Compeund ascidians
Sponge
Calcareous bryozoa
Worm tubes
Other Hydroids—
Synthecium sp.
Thyroscyphus marginatus (Bale),
Sertilaria unguicilata Busk.
Symplectoscyphus subdichotomus (Kirchenpauer ).
Halopteris campanila var. campanula (Busk),
Plurnularia proeuntbens Spencer.
Thececarpus. divaricatus var. cystifera Bale.
Halicornaria longirastris (Kirchenpauer).
Epilithic Species
Freeerowing Species
Tloldfast Species
Cc. pearsonensis 1.. anichael-sarsi
Yi Ww.
halo nara * A
Dinension, mm tyre type Africa Tndies
Stem length 1.80 1.32 = 1.00-1,50
dium. 007 017 _ O.1-0.15
Hydrotheca—
Width af margin 0.58 O68 0.25 0.25-0,30
0.69 O65 0.40 0,50
depth to diaphrasta O78
laria or Clytia. The barely discernible dia-
phragm and very delicate thecal wall suggest it
may belong to the latter genus, but its system-
atic position is indeterminate until fertile
material is found.
Campanularia ambiplica Mulder & Trebilcock,
1914: 11, figs, 2-4, Shepherd & Watson,
1970: 140.
Paracalix ambiplica (M. & T.). Stechow, 1925:
209, fig. EB.
Recards: S, 5—14 m, on red algae
Material: Three infertile stems. Stems short,
spirally annulated. Hydrothecae long, narrow,
walls parallel, a very strong S-shaped fold
about halfway along thecal wall. Margin with
6 teeth, cach with several reduplications.
Remarks: Type material of C. ambiplica in the
collection of the NMV shows more campanu-
late hydrothecue with blunter and less deeply
excavated marginal teeth than the Pearson I.
14(C), 16(C), 19(R), 20(R), 21(R), 22(R),
23(VC), 2B(VC), 29(R). 32°VE). 3R(C);
43(R), 49(C). 59(R)
15(R), 38(C), 75(R)
5(C}, 9(R). 13(R), 14(C), 15(R), 25¢R),
27(C), 28(VC), 38(C), 49°VC), 56(C),
§9(R), 74(C), 72(R)
3(R), 6(R), 10(R). 14(G), 27(C), 2B(VCd,
29(R}, 30(C), 32(VC), 35(R), 38(C),
40(R). 59(R), 75(R). BLEVC)
19(R), 59(R)
16(C)
67(C)
20(R), 56(C)
TT(RY
32(C)
16(C), 62(C)
3(R), B(VC), 19(R), 22(R), 29(R), 38(C),
4i(C)
8(VC), 17(R), 41(C), 520VC)
1(R). 58(R), 64(C), 7R(C), BOR), BICVC).
82(R), 83(C)
S7(C), 65 (VC)
2(@), 11(R), 42(C), 4300), 47(C)
specimens. Stechow's figures of specimens from
Champion Bay, W. Aust. aré intermediate in
length between the two. Meussurements are
given for comparison.
Victoria
Diwensions, nm Peai3on L. Champion Bay (M. & T)
Stem diam. 0.06 0,06 0.06
Hydrotkeca—
fongth 1,52 0.42 0.34
width DAT O18 0.17
Campanularia australis Stechow, 1924: 61.
Shepherd & Watson, 1970: 140.
Orthopytis australis (Stechaw). Hirohito, 1969:
10, fig. 9.
Records: R, 34m, on the algae Pteracladia
lucie, Cysiophora brownii, and sponge.
Material: Stems variable, to 3 mm tong, longer
slems smooth, short stems annulated, Aydro-
thecae compressed, with a wide submarginal
flange and 9-10 blunily pointed teeth, in one
instance showing reduplication. Aydranih with
16—20. tentacles, Colonies fertile.
Remarks: Although widely distributed at all
depths and on a variety of substrates, the
colonies were. not luxuriant.
162
Caumpantilings
1924; 62,
guussica Stechow, 1923: 102;
FIG, 3
Record: S24 m, on calewrcous bryazoa-
Miverial: Three infertile stems, Sremes to 4 rom
long, shewing jomts where breaks have fe-
generiuled, A spherule between stem and bydro-
theea. Hydretheeae large, campanulate, 1.26-
1,32 mm deep, expanding evenly from a nar-
row base to margin, an annular diaphragm in
thecal cavity. Margin variable in diam., 0.78—
1.2 mm, with 12 deep tongue-shaped teeth
0.015-0.618 mm wide at base, 0.013-0.015
mm long, the sinus hetween of same shupe and
size as tooth. Aydranth with approx. 16 ten-
tacles surrounding a thick annular hypostome,
similar to that of Eiclenedr/um,
Remarks; The Pearson J, specimens are among
the largest specimens of Campanularia
recorded from Ausiralian waters, the only other
specimen of comparable size being “Campanu-
lavia tinera var." of Mulder & Trebileock
(19149). from Brey Creek, Vic, (see dis-
cussion below), This is a new record for S.
Aust.
Remarks on whe states of Co tinela Hineky,
1867, C, giussica, and C. australis,
Stechow (1925, p, 206) placed Mulder &
Trebileock’s vars. ‘“‘a", “b", “co” snd “a” of
C. tineta in tentative synonymy with C. peur
sica, However, the dimensions of the hydrothe-
coe calculated from Mulder & Trebilcock's
figures are somewhat greater than Lhose given
by Stechow for his specimens. Moreover, his
figure does net show the annular diaphragm
figured by Mulder & Trebilcock, a feature alo
presentin the Pearson T. specimens.
Earlier, Stechow (1924) stated that the
gonosame of CG. gaussica was unknown, yet in-
cluded in his synonymy (1923, p. 102) a
questionuble reference to the C. tincra of Bale
(1884, p. 57) from Portland, Victoria, for
which the ponutheca was figured. He later te-
ferred Bale’s species ta C, langitheca without
explanation. The Portland specimen ih the col-
lection of the NMV is undouhtedly C, australis,
nul C. gansefew.
T cjinnot agree with Rees & Thursficld
(1965, p. 94) who reterted ©. gansyica to the
synonymy of G. dincta, The latter is a very
smal) and distinctive species; although they
stated they had examined the type material of
C. iieta, they failed to ote the very con-
siderable difference in size between the two.
The definition of the Four species, C) zincrer,
C, ausrralis, C. gatssiea and C, africatia, from
IEANETTE FE. WATSON
Austratian waters rs thus somewhat confused,
Although €, tiara and C. australis are very
similar, with bilaterally symmetries! hydro-
thecae, C’, ansrralis ts considerably the larger of
the two. ©. dugfralis shows a wide choice of
substrite, being bolh epizoic and epiphytic,
whereas C. tinete appeurs to be most Frequently
assochited with the seagrass Amphibolls antarc-
tica LW, unpublished),
Wirohity (1969, p. 10) transferred C. any-
tralis to Orthapyxts on the basis of the bilateral
symmetry of the hydrotheca, but the true
gencric status of C. australiy will remain inde-
terminate until the gonosome. is found.
C. africana is a distinct species, and has
been redescribed by Millard (1966, p, 474),
The hydrotheca is of medium size and the
gonosome is known. This species. has been re-
corded from Queensland (Pennycuick 1959, pp.
169), from “rock poals and weed”,
Uniil the collection of fertile material estab-
lishes the validity of C, waswralis and C, gays-
vica, those forms from Australian waters with
very large campanulaie hydrothecae and 10-14
tongne-shaped teeth are recognized as (". guus-
sie, those with large parallel-walled hydro-
theeae and bilateral symmetry are recognized
as C. austrafis, and those similar to C. atastralis
but much smaller in size, as C, tinere. In the
gase of C_ gaitssica, it is possible that more than
One species may actually be involved,
Campanularia sp-
FIG. 4
Records; S, 17-20 m. on red atgac.
Mfarerial: Three infertile stems. Srems 0.51—
1,72 mm long, spirally annulated, annulations
sometimes indistinct, maximum width of stem,
04 mm. Aydrothecae campanulate, length
0.39 mm, widest about one third distance
up from base, this point belng marked by a
crumpled fold encircling thecal wall. Base of
hydrotheea flat, with o slight concavity, a
socket and xpherule between hydrotheca and
stem. Margin 0.22 mm in diam, with & hroad
fongue-shoped teeth 0.025 mm high; width be-
tween tecth, 0.06 mm.
Remarks; The hydrotheeac are very delicate,
Although the hydrolthecae are somewhat col-
lapsed in mounting, the fold around the proxi-
mal region of the thecgl wall is clear in all
specimens,
The specimens are undoubledly referable to
the C. ambiplica-C. pulechratheca group en-
demic ta southern Australian waters, most
resembling the latter species, However, ©,
IL¥YDROIDS 163
Reticularia annulgfa nsp, Fig, 5—Group of three hydrothecac Fig. 6,—Distal chd of
Fig. 2 Clytia (?) pearsonensis n.sp. Hydrotheca, from holotype.
Fig. 3- Campanuluria ganssica Stechow. Hydrotheca; stem with regenerated pedicel,
Fig. 4, Campanularia sp. Hydrotheca.
Figs. 5,
hydrotheca, enlarged. Drawn from holotype.
Figs. 7
6
.8. Reticularia sp. Fig. 7.—Part of colony, showing growth habit on bryozoan colony. Fig.
8 —Hydrotheca, enlarged, showing reyenerated distal end.
pulchratheca has 14 sharply pointed teeth, the
intrathecal fold is in the distal region, and
does not encircle the thecal wall as in the
present material. This may be a new species,
but the material is inadequate for determina-
tion.
Family LAFOEIDAE
Reticularia antarctica (Hartlaub, 1904). Tot-
ton, 1930: 160, fly. 17, Briggs, 1938: 26.
Eafoas antarcti¢a Hartlanb, 1904; 11, pl. 2, fig.
Filellum
1925: 214.
Recards: R, 18-33 m, on stem of Thecoeer-
pus divaricatus var. cystiferu. S. 14 m, on
stem of Halicarnaria longirostris.
antarcticum {Hlartlauh). Stechow,
Materia/: Luxuriant infertile colonies, Hydro-
thecae delicate, of variable length, arising at
various ungles from hydrorhiza so thickly that
it is difficult to determine the length of the
adnate part, but it is usually not more than
one quarter of the total length of the hydro-
theea. Margin slightly everted, often with 2-3
teduplications; occasionally an earlier redupli-
cation about halfway along hydrotheca coin-
cides with a slight flexure of the thecal wall.
Length of free part of hydrotheca, including
reduplications, 1.0-1,18 mm; diam. at margin,
0.21 mm.
Remurks: Mt is very difficult to distinguish be-
tween R. antarctica and R, serpens in the
|i
absence of coppiniac, and as the diameter of
the (hecal margin is greater than those measure-
ments given by Stechow (1925, p. 214) for R,
serpens (after Millard 1958, p. 175), the Peur-
son |, spccimens are referred to R. antarctica,
This is the first record of R. antaretica from
S. Aust. waters. (Other locality—W, Aust.)
Reticolatia annulata n.sp,
FIGS. 5. 6
Type Material una Records: Holotype NMV
G1922, microslide: G2091, preserved
Malerial, remuinder of holotype colony—sS,
L7.m, on a small calcareous bryozoa.
Description from holotype: Hydrothecae bong,
tubular, inereasing slightly in diameter disttlly,
adnate for a sroall part of length, free pari
0.15-0.18 mm, curving out from hydrorhiza,
Hydrorecae ringed throughout eatire fength
with closely und evenly spaced annular ribs,
average distance between ribs U-US mm, each
rib with shatply everted rim; unnulations on
adgauline wall not as shatply defined us those
on free wall, Margin circulur, entire, same
diam. as hydrotheca, 0.19-0.26 mm, occtasion-
ally sinucusly curved, with everted tim. Gono-
theca wbsent,
Remarks; R. annntata is closely related to both
R. antarctica {Wartlaub, 1904) and R. serpens
(Hassall, 1848) jn shape and dimensions of the
hydrotheca, but it is easily distinguished from
these species by the close thecyl rings, FR,
serrata (Clarke, 1879) Is annulated, but the
wnnulations are confined to the adnale part of
the hydrotheca, and it is 4 much smaller
species.
The rings in R, canulatea are o£ uniform size,
and the distance between them varies. litte
along the entire length of the hydrotheca. They
have developed by continuous apical reduplica-
tion during growth of the hydrotheca, the
flange of each rth being a relict margin.
Reticufaria sp.
FIGS. 7, 8
Recerd: R, [S m, colony investing stalk of
calcareous bryozona,
Materiel; One infertile cdlony. Hydrethecue
arising in groups, or singly at irregular inter-
vals, Hydrothecac from the outer hydrorhizal
tubes adnate for approximately half their
length, free part standing, out almost perpen-
dicular to hydrorhiza. Length of free part 0.45—
0.66 mm, only the orifice of those hydrothecue
more deeply embedded in the stolonic camplex
visible. Hydrothecac tubular, 0.2 mm in diam,,
IEANETTE,
E. WATSON
many with Z—3 regeneractions after breakage.
some with marginal reduplication. Cole.e—
brown.
Remarks: The general appearance of the
hydrothecae and the diameter of the margin
ure very similar to R. antarctica from Pearson
L., but the hydrothecae of the present specimens
are much shorter; the thick woody fascicled
hydrorhizal tubes further distinguish the
present material.
The twiggy appearance of the colony, iin-
parted by the shape of the host, strongly sug-
gesis the growth habit of Cryprolaria, but with-
out the regularity of arrangement of the hydro-
thecne of that genus.
This may prove to be a new species
Zygophylax antipathes (Lamarck, 1816). Rees
& Thursfield, 1965: 76.
Sermilaria antipauthes Lamarck, 1Bl6: 11S,
Campandlaria rufa Bale, 1884: 54, pl. 1, fig. 1.
Efesoneiia antipaties (Lamarck), Ritchie, 1910;
Zygophylax rufa Bole, 1914¢: 90,
FIG, 9
Records: R. 18-45 m, among holdfasts of
hrown algue on vertical walls and on the
seafloor.
Muarevieal: Three infertile colonies, the largest
12 em high, growing from a small rootstock,
Stew woody, very brittle, main stem fascicled
(2 mm thick in largest colony) the fasciculu-
tions decreasing distally along the branches.
Branches given off randomly around main
stem. some of younger branches mono-
siphonic, /lydrothecae alternate, 0,32-0,34
may deep (margin to diaphragm) arising from
an upoaphysis at 45° to stem, frequently a short
segment (a broken und regenerated pedicel of
an earlier hydrotheca). between Nycdrothecul
pedicel and apophysis, Adcauline thecal wall
convex, 0.30-0.37 mm long, usually smooth,
sometimes. a little undulated; abcauline wall
straight or slightly concave, 0,30+0,36 mm
long. Margin usually with a distinctly everted
rim 0.17 movin diam., occasionally with | re-
duplication, Diaphragm near base of hydro-
theca transverse, occasionally oblique. Nemiato-
thecae pare, only 2 seen in mounted specimens,
one given off from a hydrothecal apophysis,
the ather from a polysiphonic tube of the stem.
Colour—aeep reddish brown,
Remarks; The branches are overgrown with
algae and conipound ascidian,
Bile (1914¢) maintained the distinction
between Z. rufa and 2, antipathes on the fal-
lowing cntenas
HYDROIDS
(i) the smaller colonies of Z. rufa, the lack
of rigidity of the branches, and,
Hid L. antipathes, following Billard, shows
no distal narrowing of the hydrotheca,
nor an everted margin.
Examination of a series of microslides of Z,
ea in the collection of the NMV, show. Bule’s
material to have come from either a broken
branch, or the distal end of a very young
colony, and there ate few hydrothecae which
are not hoticably narrowed distally; some also
lack an everted margin. One branch of the
Pearson L material bas a series of hydrothecse
with almast straight walls, no eversion of the
margin, und a rather more delicate perisarc
than usual,
Regenerated hydrothecal pedicels with an
additional segment are common in the older
regions of the stems, but are net present in the
Younger branches. They are a character de-
veloped with aging of the stem, and are thus
not specifically diagnostic.
I have compared microslides of Ritchie's
“Thetis” material of Lictorelle artipathes with
Z. rufa of Bale, and find them to be identical
in all respects, except that the perisare of Z.
rufa is much more delicate than that of the
“Thetis” specimens.
Ritchie did not comment on the presence of
nematothecae (similur to those on the Pearson
I. specimens) visible in his slides. These were
apparently noted by Rees & Thursfield (1965)
who transferred the species 10 Zygopliylex
without comment.
Since the present material has features which
clearly bridge the gap between Z. rufa and Z.
antpathes, the Wwo are considered synonymous,
This is the first record of Z. aniipathes in
S. Aust. waters. (Other localities—Torres
Strait, and off Port Jackson, N.S.W.)
Family LINEOLARIIDAE
Lingokwia spinulosa Hincks, 1861: 280, pl. 8,
Shepherd & Watson, 1970: 140.
Record: S, 15 m, on the seagrass Posidenia
ruasrrialts,
Material; Numerous infertile colonies over-
Tunning the blades of the seagrass.
Remarks: L, spintlose was not found on any
other substrate at Pearson L,
Family HALECUDAR
Ophiodissa austratis (Bale, 1919).
ph renles avairaliy Bale, 1919- 396, pl. 16, fig.
Record: R, 19 mi, On black sponge.
165
Material: One colony of several infertile stems
growing from a matted hydrorhiza on the sur-
fuce of the sponge, Sremx to 2 cm long, fas-
cicled, itregularly branched, with 2-3 supple-
mentary tubes extending two thirds the distonce
up stem. Aydraphore with a few reduplica-
tions. WNematothecae rare. Crleour—light
greenish, with black patches scattered through-
out hydrocaulus. (Under the microscope these
pitches are black granules concentrated on the
hydranth and in the coenosyre.)
Remarks: The status of Ophiodissa haa been
briefly discussed by Watson (1969, p, LIN),
Bale (1919) described, but did noe figure the
gonophore of O. australis, Ralph (1953, p,
342) way. uncertain whether @. australis is a
synonym of Hydrodendroy — cacinifarmis
(Ritchie, 1907) but kept the two apecies
separate because “the hydrothecae of 4, aus-
tralix Bule are shallower, measured fromm the
margin to puncta line, than those of H. cuvini-
formis, und the goncthecae of the latter are
unknown". Millard (1966b, p. 490) described
the gonophore of H, caciniformis fiom material
from the Vema Seamount. off the west coast
of South Africa.
There are two microslides of O. australis in
the Bale collection In NMV, one trom Green
Point, N.S.W,, and the other from Port Phillip
Heads. Vic. The latter specimen is 4 lightly
fascicled stem 2 cm long, with u group of 10
male gonophores growing {rom the hydrohiza,
and is undoubtedly the slide fram which Bale
described the gonotheca of the species.
The gonothecae aye smooth or very slightly
wnnulated, with curved or straight pedicels, and
several have a slight constriction just below ihe
truncated distal end, The gonopheres are
nearly mature, the blastostyle almest filling the
gonothecal cavity, aod above the blasroscyle
there ts a ring of black granules,
The gonothecae and gonophores of Bale’s
material are similar to those of A, caciniformis
figured hy Millard, but the ponotheca of O..
australis is much longer and more than twice
the width of those of JT. caciniforinis, Further-
more, the hydraphote of HW. cacinifprmiy (trom
Millard’s figure) is both wider and deeper than
that of Q. ausiralixs from Green Point, It seems
that the two species, while very similar, are
distinct,
Comparison of measurements (see helaw)
of HH, caciniformis from New Zealand with
Bale's Q. australfe shows that the New Zea-
land maternal Falls near the dimensional range
of OO. aeuwealls, and may well be this species.
Lob JEASETTE, E, WATSON
The finding of fertile material in New Zealand
waters will settle this. point.
O. australis
Dimensions, mm H. caciniforialy
Pearson Grein N: 8.
I, Pi. Fealand = Africa
Uycirophore— .
diam, 1 Paneta dine 0.14 0.15 0,12 0.20
depth, margin to
puacta ne 0.025 O05 0.44-4,06 0.10
Gonotheca
length t.441.50 — 6 90-1,90
width at mporpyre — 0.94-1.00 —_ 0.40
The specimens from Pearson I. are identical
im every respect with Bale’s 0. australis. This
is the first record of O. awsrralis from §. Aust.
Ophiodissa blackburni a.sp.
FIGS, 10-12
Type Material and Records: Hoaletype.
NMV G1927, microslide; G2092, preserved
material, remuinder of holotype colony—-S,
27 m, on Herdinuiia monus; Paratypes
GI928, G1929, microslides; G2093, pre-
served material, reémuinder of paralype
colony—S, 24-27 m, on bryozoa and
sponge.
Description from holatype ard paratypes:
Hydrorhiza a winding tubular stolon 0,11-0,13
mm in diam,, thick and strongly corrugated
throughout entire length, becoming erect at
intervals 10 form monosiphonic stems. to 12
mm high, Uydrephores given off irregularly,
either directly from the siolon or from stem,
Pediccl of hydrophores of variable length,
0.19-0,27 mm, beginning with an annular con-
striction 0,07-0,10 mm in diam., followed by
1-2 annulations, then expanding evenly to: mar-
gin, Perisarc thin. Secondary and tertinr'y
hydrophores common, branching outwards just
below the diaphragm ot primary (or secon-
dary) hydrophore, becoming ascending at an-
nular constriction. Hydrophores: redupliculed
up to 4 times; reduplications of variable length,
given off successively from the diaphragm of
preceding hydrophore, Diaphragm 0.13-0.17
mim in diam., moderately deep, 0.04 mm from
margin (best seen in preserved material). Afur-
vin flaring, with strongly everted lip, diam,
0.22-0.24 mm. Neutothecae sparse low tubu-
lar orifices 0.03 mm high, and 0,05 mm in
diam, at base, situated on hydrorhiza or stem,
opposite, or nearly opposite hydrothecal pedi-
ecl. Aydranth large, extensile, with approx. 30
stubby tentacles. Co/lawr—yellow, Gonetheca—
absent.
Remarks; The presence of nematothecae places
the present material in Ophiodissa (Watson
1969, p. 111). Although in. some instances the
hematothecae may be mistaken for the broken
hase Of a hydrothecul pedicel, their position
opposite the pedicel, and their smaller size
usually serves to distinguish them.
O. blackburni shows some resemblance to O.
carrugata Praser, 1936, However, neither
branching nor reduplication of the margin is
mentioned or fizured by Fraser, and the ten-
ticular orguns of O. currugata are described as
being relatively large and flaring slightly at the
marfgin (Fraser 1936, p, 113).
Blackburn (1938, p, 322) described a frag-
mentary Haleciunt sp. Crom the Jittoral zone
of Reeyesby L in the Sir Joscph Banks Group.
S. Aust., remarking that the material, too ob-
scured by forcign matter for diagnosis, Was
probably a new species, similar to A. corra-
garuen Nutting, 1912. Although Hlackburn's
specimens are not available for comparison, it
is certain that his milerial aimel the present
specimens are the same species.
Phylactotheca armata Stechow, 1924: 54:
1925; 204, fiz. C. Blackhurn, (942° LOA,
Ophtiodiata Jrogitts Blackburn, 1937a! 365, fig.
Records; R, 33 m, on sponge and ascidian,
Maserial: A few infertile stems to | em high.
Renuirks; The Pearson I. material conforms to
the description of the species given by Stechow
and Blackburn.
There are no secondary hydrophores, nor
any nematothecac developed in the present
material, Blackburn's specimens from Balnar-
ring. Vie., similarly showed no sign of nemu-
iothecac, Hence their presence or absence cam
not be taken as a good diagnostic character
lor Lhe aenus,
Halecium delicatulum Coughtrey, 1876: 299,
Ralph, 1958: 334, figs. 11, 12. (syno-
nymy),
Records: R, 21-45 m, on Plamularia pro-
cemberns and Syntheciunt sp.; 8, 24 m, on
Herdmania momus, and brvozoa,
Material: Miny small infertile stems to 2 om
high. Sresns irregularly and sparsely branched,
a few are lightly fascicled, with an extra poly-
siphonic tube running up the proximal part of
the stem.
Renuirkay Oye of the mest abundant epizoic
species tm the collection,
HYDROIDS UG7
Halecium sp. 1.
FIG, 13
Record: R, 30 m, an lower stem of Halj-
cernarie fongirostris,
Moreelul’ A single infertile stem L2 mm high.
Stem unfascicled, irregularly branched; stem
and branch intemodes of vanable length, up
to 0.22 mm, and 6,04 mm in diam., annulated
proximally, but otherwise fairly amooth-
Branches given off from distal end of inter-
nodes on lower part of stem, Pedicel of pri-
mary hydrephore of yariable length, proximal
node transverse, follawed by 1-2 antulations,
the remainder smooth, expanding evenly to
diaphragm. Aydrophere small, shallow, with
circular margin, 0.08-0.12.mm in diam., and
strongly everted tim; up to 3 reduplications
given off successively from mouth of preceding
hydrophore. Diaphragm present, depth from
margin to diaphragm, 0.02-0.03 mm. Secearn-
dary and tertiary branches comman, arising
from distal end of pedicel of primary (or
secondary) hydrophore,
Remarks: This very small form resembles H.
tenelliom Hineks, 184) in size and delicacy of
the trophasome, btit the dimensions are ever
smalfer than those given by Millard (1957, p.
193) and Rilph (1958, p, 340) for this
species.
in the present material, only the first inter-
node can be described as a true stem, the
branching being truly arborescent, all sub-
sequent internodes being secondary branches
given off the pedicels of the primary hydro-
phores. The presence or absence of punctae
cannot be ascertained because of foreign mat-
ter, However, the Bale collection in the NMV
contains a microslide labelled “Halecium, Grif-
fiths Point, July, 1880" with one small infertile
stem identical with the Pearson J, specimen,
and the hydrophores of this stem show « very
clear rung of punctae between the margin and
diaphragm.
This is probably a new species, but due to
ils similarity to HW. ¢enellam its identity is in-
determinate until adequate Fertile material is
found.
Halecium sp. 2.
FIG. 14
Record: R, 35 m, on algae,
Murerial> Two yery small infertile stems.
Aydrerhize tubular. Stents unbranched, 2 mm
long, internodes of variable length, 0,28-0.66
rom, dian U.08-0,12 mm, perisare thick,
divided into segments by 2-3 deep, transverse
constrictions, the last segment expanding
slightly to support base of hydrophore. Hydre-
phore very shallow, depth to diaphragm 0,03
mm. Margin circular, 0.14-0:17 mm in diam.,
slishtly everted. Succeeding stem internodes
arising just below hydrophore, standing out
perpendictilarly, giving siem a zig zug appear-
ance. Hydranth too large to retract into hydro-
phore, body thick, about 24 stubby tentacles,
Remarks: All the bydrathecal margins are so
damaged that it is impossible to determine if
punctae are present, Although the dimensions
fall well within the range of J. lartkesteri
(Bourne, 1890) (Millard 1968, p, 257) the
internodes show no tendency to curve upward,
nor is there any sign of the secondary hydro-
phores common in this species. For these rea-
sons, the present specimens are not assizned to
A, fankestert.
Family SYNTHECIIDAE
Svyothecium elegans forma subveniricasum
Bale, 7914, Ralph, 1958: 347, fis. 16,
FIGS, 15. 16
Synthecium elegans Allman, ($72: 229, Black-
hurn, 1942; 111.
Synthecium subyentricosum Bale, 19)4az 5, pl.
1, figs. 3-5; 1915: 265,
Records; R, 34 m, on Herdmenia memeasy
S, 18-25 m, on stem of Therocerpuy eivuri-
catus var. eystifera; Stn. F, 65 m,. on worm
tubes with Diphasia subcarinate
Marerial: Several immature colonies: one fer-
tile stem. Stems monosiphonic, flexuons, to 2
cm long; no secondary branching, but a few
tendrils given off distal ends of hranches, Proxi-
mal stem internodes with 1-3 pairs of opposite
hydrothecae; succeeding internodes 1,5-1.8
mm long, a pair of opposite hydrothecae in
middie of internode, und u pair of opposite,
distally situated hydrocladia. Aydrothecae of
variable shape, generally tubular. three
quarters of length adnate to intemode, free
adcauline wall 0,07-0.15 mm Jong, fixed wall
0.42-0.45 ram: abcauline wall 0.58-0.44 mm
long. A delicate internal sheath clearly visible
i many hydrothecae, Murgin of hydrutheca
sinuous, sligbtly everted, O18-0.19 mm in
diam., a few marginal rcduplications. Gero-
theca—a single immature individual prowing
from the orifice of one of the second pair of
hydrothecue on basal stem internode. Calour
—white, with trace of purple.
Remarks: The single immature gonothecs iden-
ties the Pearson J, material with 8, swhvenrri-
casum Bale. recognised by Ralph (1958) as a
varietal form of §. elegans Allman,
168
JEANETTE BE, WATSON
15
Zygophylax antipathes (Lamarck). Portion of branch showing hydrothecae with everted
margins and regenerated hydrothecal pedicels,
. Ophiodissa blackburni n.sp. From holotype. Fig, 10—Hydrorhiza and stem with secon-
dary hydrophorcs and reduplicated margins. Fig. 11.—Reduplicated hydrophores, en-
larged Fig. 12.— Nematothecae, enlarged.
Halecium sp, 1. Whole stem, showing growth habit.
Halecium sp, 2. Part of stem showing hydraphores and growth habit.
» Synthecium elegans forma subventricosum Bale. Fig. 15—Part of stem. Fig. 16—Imma-
ture gonotheca.
. Synthecium dentigerum Jarvis, Fig. 17W—Part of branch with subupposite to opposite
hydrothecae. Fig. 18—Hydrothecae enlarged, showing internal adcauline teeth.
HYDROIDS 149
Although there is considerable variation in
shape of the hydrothecac among the specimens,
most aré tubular and closely adnate to the
hydrocladium, not ventricose, as described by
Bale for his specimens from the Great Austsa-
lian Bight, Morcoyer, there is only one pair of
opposite hydrothecae in the middle of each of
the distal stem internodes, and a pair of oppo-
site. distal hydrocladia, and the hydrothecal
marvins are decidedly sinuated. all features
considered by Bale to be criteria distinguishing
S, parulum Busk.
Thus, S. vlegany forma subventricasiuint may
eventually prove to be a synonym of 3. pata
lum. However, in view of the lack of adequate
fertile material, the fact that the typo of &S.
patulernt is not known to exist, and the possi-
bility that sexual dimorphism may occur, it is
hest to keep the species separate at present.
Syutheciurt dentigerum Jarvis, 1922: 344, pl,
25. fig. 15. Totton, 1930: 172. Millard,
1944-24. fig. 6,
FIGS. 17, 18
Records: S, 23 mon stem of Sertnlaria un-
gaiculata and Meramenntig momus
Marerials Five infertile stems, to 2 em long.
Stas monosiphonic, basal internodes fong..
3443.6 mm, with 5 pairs of opposite hydro-
thecac: each succeeding internode with 2-3
pairs of hydrothecae in mid region, followed
by a pair of opposite hydrocladia. Aydrocladix
ansing perpendicular to stem from a distinct
proximal joint; unustomoses. and some secon-
dary branching oceur, Aydrorthecue almost
titbular, adnate for two thirds their length,
fixed adcauline wall 0.45-0.51 mm long. free
wall (.15-1.2 mm: abcauline wall 0,36—-0.42
mm. Murgin narrow. sinuated, with slightly
everted cim, 0.21-0.27 mro in diam. Proximal
hydrocladial hydrothecae subalternate, hecom-
ing opposite distally. {One stem has alternate
hydrothecae with the abcauling wall hent
sharply outwards.) Adcauline submarginal
tooth present in most hydrothecae, variable
from wedge-shaped to a mere thickening of the
udeauline wall, No ahcauline teeth or marginal
teduplications,
Remarks: Three specics of Synthecine with
internal teeth have been described. §. carinassei
Totton, 1930, 8S. singulare Billard, 1935, and 8.
dentigeriuim Jarvis, 1922. The Pearson [. speci-
mens are larger than 8, carinatum and smaller
than 3. wingulere, but fall well within the range
giver by Millard (1964) for S. denligernen.
The specimens are distinguishable from S$. ev.
guns forma sithventricosum Bale, from Pear-
son 1, only by the presence of the internal suts-
marginal tooth, which is however not
developed at all in some hydrothecae, the most
prominent tecth being associated with those
hydrothecae which have a flexure of the ab-
cantine wall, Although the present matertal
differs in. some respecis [rom desenptions of §,
dentigerym in size of the colonies and arrange-
ment of dhe stem hydrothecne, these are neither
sulfigicntly constant nor importam characters
to Wattant the erection of a new species,
This is the first record of S. dentiveruen from
Austrahan waters. It has been reported twice
previously, from the Indian Ocean pnd South
Africa.
Fumily SERTULARRDAK
Thyroscyphus marginatus (Bale, 1884), Bale,
1915; 245, Stechow. 1925: 217. Black-
burn, 1942: 112.
Campianularia marginata Pale, 1884; 154, pl. 1,
fig, 2; 1888; 758: 1914) SI. Bartlelt, 1907: 42.
Records: R, [4-30 m on algac and sponge;
5, 22-30 m on Sergewsum varians
Material Four infertile colonies ot u few stems
each. Aydrorhiza a simple tube, lagsely wound
on substrate. Sven simple, to 1 cm lung,
smooth or slightly anoulated, diam. increasing
distally, a distinct transverse joint just shove
junction with hydrorhiza, Pedicely of hydro-
thecac 1.95-2.7 mm long, 0.25 mm in «liam
distally. In one stem, 2 branches ate given off
side by siule, 1 short, terminating in a hydro-
theca, the other continuing normal growth. giv-
ing off a pedicel distally. Hydrothecal marain
with +4 teeth, 4 valved operculum, and
thickened submarginal ring, Depth of hydro-
theca, 1.5 mm: diam. at margm, (.73-1-21
min.
Remarks: This agrees fairty well with descrip-
tions of T. marginatus given by Bale.
Parascyphus simplex (Lumouroux, 1816).
Blackburn, 1942: 112, Ralph, 1961a; 755,
fiz. 1. Stechow, 1925; 224.
Laomedea simplex Lamouroux, 1816; 204.
Cumpanularia siraples (Lamouroux). Bale,
1884: 58.
Pantponaitaris raridentata Hale. 1894" 98, pl, 3,
&, 3,
Tivrascyphus simplex CLameureux), Hodgson.
1980; 10, fiz, 22.
Records; R, 34 m, on sponge and Aere-
maria moms on vertical walls; § 25 m, an
Thecocarpus divaricarns,
Murerial) A few infertile colonies, Sreme short.
unbranched, t9 5S mm long. Averothecae
170
swollen on proximal adcauline walls same with
a short pedicel, Stem apophyses pronounced,
many with a constriction marking the site of
growth regeneration after breakage.
Remarks: The stem apophyses figured by Hudy-
son (1950) and Ralph (1961) appear to merge
into the base of the hydrothecae, Bale’s (1894)
figure shows a more pronounced apophysis.
similar to those of the Pearson TI. specimens.
Diphasia subcarinata (Busk, 1852). Bale, 1884:
102, pl. 4, fig, 1. pl. 19, fig. 18; 19144; 7;
1915; 264, Ritchie, 1911; 850. Hodgson,
1950: 20, figs. 34. 35. Ralph. 1961a: 764.
fig. 5, Shepherd & Watson, 1970: 140.
Serratia sithearinata Busk, 1852 390,
Records; R, 20-34 m on bryozoa; 8, 25 m
on Amansia pinnatifida and Herdimania
mranuiss Str. F635 m, on worn tube.
Maretialy Many infertile unbranched colonies
0.5-1 em high. C'oleur—dark brown.
Remurks: ‘The keel deseribed by Bale (1884)
in his Victorian material is ptesent in the Pear-
son [. specimens only as an indistinct ridge
passing between the Jateral maryinal tooth and
the adewuline thecal wall. Ralph (1961) notes
the presence of the ridge as a “prominent fea-
ture” un her New Zealend specimens, but
Hodgson (1950) was unable to find the ridge
in his. Tasmanian material.
Stercatheca elongats (Lamouroux, 1816).
eae Lene, 23. figs. 38, 39, Ralph,
1961: 762, fig. 4. Shepherd & Watson.
1970; tag
bert itaria sian Lamouroux, 7816: 189, pl,
Bale, 1884; 75, pl, 6, figs. 7, 8 pl 19, re,
7: 19)3: 277. Mulder & Trebitcock, 19t4a; 3,
pl, t, figs, 7-10.
Records: R, 30-45 m; S, 12 m, of Srene-
claudia australis, Phacelocurpus lahillaradiers,
Scvrathalia dorvearpa and Aerovarpia pani-
culdta.
Materiel: luxuriant colonies, some fertile.
Sremy 4-5 cm long, Gonothecae with long
horned processes.
Remarks: §. clangata is one of the commanest
epiphytic hydroids of the southern Australian
coastlins. The Pearson f. specimens correspond
to the “short stemmed ocean form" of Mulder
& Trebileoek (19148). 8. eloneuta shows a wide
choice of algal substrate and consideruble
tolerance of environmental conditions. The
stems of the present specimens Were fice of the
encrusting coralline alga usually axsocinted with
this species,
IFANETTE k, WATSON
Crateritheca sacanthostoma (Bale, 1882),
Ralph, !96la; 756, fig. 2. Shepherd &
Watson, 1970: 140. Millard, 1964; 26,
fig, 7.
Sermlaria ariel embeeeis rn 1882; 23, pl £2.
Fig. 4; R84: 8&5, pl. 4, figs. 7. 8: 1973: 434.
Bartlert 1907; 44, hile * Trebileock, 1914)
Stereotheed acanthesionta (Bale), Stechew:
1919; 103. Blackburn, 1942. 112.
Records; R, 24-33 mm. of sponge and red
algae,
Material: Two colonies of 4 few stems each.
Stenisto 32cm long,
Remarks: The Specimens agree exactly with
Bale’s description of 3, acanthosrania.
Crateritheca crenata (Bale. 1854),
1941a: 757, fig, 2.
FIG. 19
Sertalaria crenata Bale, 1884: 86, pl. 4, fig. 2.
Record: R, 33 m, on brown algue.
Material: Two fragmentary infertile stems.
Hydrorhiza tubular, winding. Sten fragments
2.em Jong, with 4-6 deep proximal annulations,
followed by a short athecate internode below
first branch, Internodes 0.13-0.26 mm_ tong.
0.04-0,06 mm in diam. ai node.
Remarks: The Pearson 1. specimens have a
much less pronounced outward bend of the dis-
tal thecal wall than C. crenara figured by Bale
(1884), However, a microslide of C. cremara
trom Snapper Point, Vic. in the Bale collection
of the NMY, compares very closely with the
Pearson [. material.
This is the second record of the species, and
4 new record for §, Aust. (Other locality—
Port Phillip Bay. Vic.)
Salacta obliquanoda Mulder & ‘frebilock,
1914).
Ralph,
FIG. 20
Sertularia obliqtanoda Mulder & Trebilcock,
(9)3b: 41, pl. 5, fig, 1, Stechow, 1924; 106.
Shepherd & Watson, 1970; 140),
Records: R, 18-46 m, on several species of
red algae, including Metunnstophora flabel-
leva, bryozoan, und sponge,
Meverial; Luxuriont colonies. — Epiphytic
colonies fertile, Aydrerhiza loose, tubular.
Stems simple, to 8 mm high, some giving olf
distal tendrils which form new stolons, Stem
Interrodes variuble in length, 0.54-0.46 mm.
width at base of hydrotheca, 0.18-0.24 mm.
joints oblique, best developed on short inter-
nodes, indistinct or absent on long internodes,
Hiydrowivese similar to the description given
ELYDROIDS Wt
by Mulder & Trebilcock, but the hydrothecal
aperture is oblique, mot vertical, sloping
diagonally back towards the stem in a line:
parallel with the abcuufine wall. Length of free
adcauline wall 0,12-0.18 mm, fixed adcauline
wall, 6.24-0,27 mm, abcauline wall, 0.21-0.30
mm long. Margin with 2 shatp adcauline teeth,
and q thickening of thecal wail in the base ot
the abcuuline sinus; width of margin 0.06-0.15
mm. Gonothecae large, barrel-shaped, |-53—
1.6 mm Jong, 0.96-1.06 mm wide, with 6-8
deep annulations, a circlet of hooked teeth be-
law tim. Usually 1 gonotheca on a stem, borne
helow the proximal hydrotheca; male and fe
male gonophores on the one colony,
Remarns; With the exception of the somewhat
longer stem internodes and more oblique aper-
ture, the Pearson I, specimens compare closely
with the type of Sertu/uria ebliqvaneda in the
collection of the NMY.
Blackburn (1938, p. 31% 1942. p. 113)
listed Dynamena cornicing McGrady, 1858,
among the hydroids of the Sir Joseph Banks
Group, D. cornicina (=Serrularia complexa
Clarke, 1879) (see Bale 1885, p. 769; Billard
1925, p. 1838) and S& obliquanods are very
similar, and are difficult to distinguish oxeept
in fresh fertile material. However, the hydro-
thecae of D. cornivine ave larger, and the aper-
tural teeth are more Jalerally situated, The ab-
cauline opercular flap, easily visible in the
present material. clearly distinguishes 5. obfi-
quanode frum D. carnicine.
The present apecimens also resemble Triden-
tara rurbinata ({Lamouroux, 1816) (Stechow
1925, p. 223) particularly in the presence of
the abcauline fiap ond the thickening of the
abcauline wall. ‘The status of this group needs
further elucidation,
S. obliqnanade is one of the commonest
species in the collection, This is the first
definite record of S, ebliavanada for S. Aust,
(Other localities—Torguay and Barwon Heads,
Vic, }
Sertularella robusta Coughtrey, 1876; 300, fig.
22: Blackburn, 1942; 115. Hodgson, 1950:
33, fig. 53. Ralph, 19la: 824. fig. 22,
Shepherd & Watson, 1970: 140.
FIG, 71
Records; R, 24-33 m: S, 15-24 m, on Ballia
callitricha, Laurencia elata. Herdmania
momus, bryozou. and sponge,
Marerial> Numerous infertile colonics, Stems
simple, iinbranched, te | cm Jong, arising fram
a tubular fydrerhiza, Stern internodes variable
in length, O.42-1,14 mm, but fairly constant in
moimum width, 0O15-0.18 mm. measured
just below hydrotheca. Aydrothecae distal on
jong internodes, occupying most of the length
of short internodes. Length of free adcauline
wall 0.3 mo, fixed adcauline wall 0.24 mm;
abcauline wall 0.42 mm; maximum width af
hydrothees, 0.24 mm. Thecal walls moderately
to faintly annulated with 2-3 broad undulations
passing around widest purt of hydrotheca.
Remarks: Following Ralph (1961a), Lhose
stems with a fairly thick perisare aid annulated
thecal walls, even though the unnulations may
be Faint, are assigned to S. robaester.
Sertularella simplex (Hutton, 1873). Ralph,
19fla; $21, fig, 21.
FIG. 22
Sevtularia simplex Hution, 1873; 257.
Sertulerella peregeina Rule, 1926; 19. fig, 4.
Records: R, 30 m, on the stem of Thece-
carpus divaricatus var, cystifera, Herdmania
momus, and on bryozoa.
Material: Colonies of a few Fertile stems, Streams
to 1 em long, occasionally branched, arising
from a tubular hydrorhiza. Stems smouth,
proximal internode athecate, with a few in-
definite snnulations, nodes distinct, sloping al-
ternately right and left, Internodes fairly tong,
0.39-0.60 mm, becoming progressively shorter
distally, until hydrotheca occupies about two-
thirds of internode. Width of internode below
hydrotheca, 017-020 mm. Branches, when
present, arising just below the hydrotheca, the
first branch internode with 2-3 annutations.
Tendrils present, growing from the distal ends
of stems and from broken hydrathecae. Hydro-
thecae with 3 internal submuarginal teeth—l
#beauline, 2 adcauline; thecal walls thin and
smooth, length of free adeauline wall variable,
0.33-0.40 mm, fixed wall, 0.25-0.27 mm;
abcauline wall, 0.52-0,54 mm: maximum width
of hydrotheca, 0.22-0.27 mm. Margin showing
occasional reduplications,
Remarks; The material from Pearson I,,,
although variable, falls well within the cange
of variation of S. simplex defined by Ralph
(1961a). The specimens assigned here to S.
simplex are easily distinguished from 4, rehbuxia
from Peurson 1, by the generally larger and
smoother walled hydrothecae-
Bule (1926) erected Sertnfarella peregrina
to include hydroids from Bass Strait and Port
Phillip formerly referred to Sertularella paly-
zonias und S. guudichardi. S. peregrina is indis-
inguishable from §. simplex as now defined,
72 JEANETTE
accordingly S. peregrina is here referred te the
synonymy of S. simplex.
A new record for §. Ausr.
Sertularellu annulaventricosa Mulder & ‘Trebil-
cock, 1915: 34, pl, 7, fig 1, pb 8, fig. 4.
FIG. 23
Senilareta undulata Bale, 1915: 284, pl, 46,
Hig. ). Hodgson, 1950; 34, fig, 59,
Records: R, 33 m, on Sarpaysunt sp., on red
algae, und bryozoa.
Muterials Colonies moderately abundant, infer-
tilt, Hydrorhize wubular, Siems simple, to 5mm
long, with 2-4 hydrothecae (exceptionally, i
stem has 10 hydrotheeae; another is branched)
hut. many hydrotheeve arising singly from
hydrorhiza, Stems annulated proximally, annu-
lafions extending to buse of first hydrotheca,
{nternedes 0.30-0,51 mm long, width helow
hydrotheca, 0.18 mm. /lydroecaue barrel-
shaped, 0,27-0,36 mm in diam, at widest part,
Margin 0.15-0.24 mm in diam., depth trom
margin to base, 0.30-0.36 mm.
Remarks: The holotype microslide of S. annti-
laveninicesa in the collection of the NMV has
longer stem internodes than the Pearson I.
specimens, and is unbranched, although Mulder
& Vrebilcock note that “one specimen shows
signs of having been slightly branched”. Vhe
hydrothecal walls of the type are fairly. smooth.
with w ledge passing around the hydrotheca u
little below the margin. The walls uf the Pear-
sou 1. specimens are not a3 smooth as these af
the type, and the submarginal ledge is replaced
by I-2 annular ridges. giving the hydrotheca a
crumpled appearance. As in the type material,
the aperture of the present material is some-
what variable in diam., ranging from 4 narrow
orifice ta almost the complete width of the
hydrotheca-
‘This is the third record of the species, and
a new récord fur S. Aust. (Other loculities
central Victorian coastline, ancl Tasmania).
EF. WAISON
Sertulurella ayrilia n.sp.
FIGS, 24, 25
Type Maternal and Records: Holotype, NMV
GI%64, microslide; paratypes, Gi96S,
G1966, G1967, inicroslides; SAM H35,
microslide; holotype and paratypes from 8,
13 m, On Sergessunr verruculosum.
Dexerigtion from holotype ana puraly pes: Stems
simple, short, to 4 mm Jong, unbranched, aris-
ing from a thin tubular hydrorhiza. First stem
intetnodes short, with 3 proximal annulations:
succeeding internades of variable length, 6.32—
0.53 mm, widening from w narrow oblique
proximal node, 0.04-0,.08 mm in diam, te base
of hydroiheca. Nodes sloping alternately left
and right, with I-2 oblique annulations, re-
mainder of internode smooth, Mydrothecae dis-
tal on internode, alternate, a. maximum of &
on stem, each sloping outwards parallel with
line of internode, giving stem a zig zaz appear-
ance. Body of hydrotheca long, almost. cylin-
drical, slightly swollen at junction with inter-
node. 016-019 mm in diam. Adcauline wall
with 5—7 uniform, rounded annulations, most
prominent across widest part of hydrotheca,
but reduced to 4-6 on abcauline wall, fading
out proximally. Adcauline wall arched out-
wards, abcauline wall inflexed into .a long, nar-
tuw neck, 0.10-0.20 mm at narrowest dian,
expanding again to margin. Length of free ad-
cauline wall, 0.36~0.45 mm, fixed adeauline
wall. 0.12~0.14 mm; abcauline wall, 0.32-0.37
mm. Aperture facing outwards and slightly
down. Margin, 0.11-0.14 mm in dian, with
4 broad, low teeth, the adcauline tooth most
prominent; 3 strongly developed internal sub-
marginal teeth in theeal neck—2 identical
dorso-lateral bract-like teeth, and | long peg-
shaped sbcauline tooth. Ayedranth — in-
sufficiently preserved for description, but shows
evidence of an abecauline caecum. Gonethecu—
1 immituce individual, arising from the base
SK SSeS
One stem jnternode showing hydrothecac
Fig. 19. Craterithecu crenata (Bale). Part of branch with three hydrothecae.
Fig- 2d, Salieia ebliqnanoda (Mulder & ‘Trebilgock),
with abcuuline opercular flap.
Fig. il Serlaretla robusta Coughtrey. Part of stem showing hydrothecae with shallowly anniu-
lated walls, and hydranth with abcauline caecum,
Fig. 22. Seetularclla sitnplex (Wutton). Part of stem showing smooth-walled hydrothecse:
Fig, 24, Sertularella annulaventricosa Mulder & Trebileock, Pact of stem.
Figs. 24.25. Sertularella avrilia m3p. From holotype, Fig, 24.—Whole stem. Fig, 25,—Part of stem,
enlarged, showing hydrothecac and internal submarginal teeth,
Fig, 26, Seriularetia sp. 1, Whole stem with two hydrothecae,
Fig. 27. Sermlaretla sp. 2. Part of stem with two hydrothecue,
Figs, 28-90. Symplectoyesphus vostratus tsp. Big. 28.-
Stem with one hydrotheca and ponotheca,
from holotype colony. Fig. 29.—Stem with two hydrothecae: from paratypes, Fig. 30-—
Hydrotheca, lateral view, showing internal submarginal teeth, from paratype.
HYDROIDS 173
174
of a proximal hydrotheca; body strongly ribbed,
with signs of the development of a slender ter-
minal neck and 4 blunt spines.
Remurks: &, aveilia resembles both 5. roluse
(Couhtrey, 1876) and S. gilehristi Millard,
1964. However, 3. aveifia is smaller than S.
robusta, the hydrothecae jee more strongly
anuulated, and it is further distinguished by the
long, curved thecal neck, Although S. evrilia
shows some affinitics with S. gilchrisri, this
Seuth African species is a larve, branching
form, with fascicled stem, and a shorter, Icss
conspicuously arched thecal neck,
Sertulurella sp, 1,
FIG. 26
Revords: R. 21-33 m, on Lawreneia elata,
Herdmania momus, Halopteris campanuia
var. campattula, and bryozoa.
Material; Numerous infertile colonies, Stems
simple, monesiphonic, unbranched. straggling.
arising from a tubular hydrorhiza 6,05 mm in
diam, Stems 10 3 mm long, first stem internode
short, with 2-3 indefinite annulations, following
internodes thecate, of variable length, O0.06—-
0.24 mm, irregularly annulated except just be-
low hydrotheca, Nodes distinct, sloping alter
nately right and Jeft, 0.04-0.06 mm in dian,
measured just below hydrotheca, Aydrothecae
distal on internode, a maximum of 4 on stem:
body of tfydrotheca barrel-shaped, one third to
one half free of internode: Jength of fixed ad-
cavline wall, 0.11-0.14 mm, free adcauline
wall, O.15-0,24 mm) abcauline will, 0,25-0,34
mm; maximum width of hydrotheca at junction
of udeuuline wall with internode, 0.20 mm,
hydrotheca gradually narrowing to margin.
‘Thecal wall with S—7 strong, entire annulations,
most marked in mid-region, annulations fading
out towards margin, Margin variable in diam.,
0,08-0.13 mm, with 4 sharp, equidistant tecth,
Nydranth with abcauline caecum, Colaur—
yellow-brown.
Remarks; The species resembles §. angulosa
Bale, 1894 (= S. robuyta Ralph, 1961) but is
a smaller species, with much more deeply in-
cised thecal ridges. [t is further distinguished
from §. rebusta by its straggling growth habit
and the large oumber of hydrothecse arising
directly from the hydrorhiza.
Although displaying » wide choice of sub-
sttate, occurring on algae, ascidigns, hryozoa,
und other hydroids, this Sertutdrelld was
recorded only on the exposed site, where is was
very abundant.
JIEANKTTE B, WAISON
lt seems likely that this is 2 new species,
tut is indeterminate until fertile material is
found,
Sertularetla sp. 2.
FIG. 27
Record: R, 18m, no substrate recorded.
Marerial; Threc fragmentary infertile stems de-
tuched from the hydrorhiza, Stems straight, un-
branched, occasionally bent slightly at u node,
pensare very thick und brittle, Intemodes of
variable length, 0.48-0.96 mm, nodes oblique.
sluping alternately lefrand right, angle of slope
variable, Internode narrowest at node, 0.18-
0.24 mm in diam., followed by 1-2 proximal
annulations, widening to base of hydrotheca;
width of internode at base of hydrotheca, 0.27—
0.36 mm. Hydrothecae occupying distal half
to two-thirds of internode, sfanding out at about
30)° to axis, very large, variable in shape, either
cylindrical, or with a slight distal narrowing be-
hind margin, 5 distinct annulations passing
completely around hydrotheca, the strongest in
the mid-region of the abcauline wall, fading
out proximally, annulations Jess distinct on
alder hydrothecue, Length of free adcauline
wall. 0.39-0.48 mm, fixed adcauline wall 0.36—
0.45 mm; abcauline wall 0.66—-0.72 mm. Mar-
gin O.33--0.51 mm in diam.. thickened, with 4+
low, blunt, slightly everted teeth. Operculum
of 4 flaps. Hydrenth not well preserved, but
an abcauline caccum may be present.
Remarks: The hydrothecae are distinctive in
shape, and are the largest recorded in Austra-
lian Waters for Sertularella. This is almost cer-
tainly a new species, but confirmation nist
awail the collection of adequate and fertile
material.
Symplectoscyphos Jongithecus (Bale, 1883).
Sertularella longithecu Bule, 1888: 762, pl. 16,
fie. 5, 6; 1894: 101, pl 4, figs 7-9.
Record: Sta, F, 65 m, no substrate recorded.
Murertal: A few straggling infertile branched
stems to ¥ cm Jong. Stem internodes straight.
0.54-0.75 mm long, nodes well defined, slop-
ing alternately right and Jeft. Branches given
off from front of stem opposite base of hydro-
thecu. Hydrothecue Jong, tubular, narrowing
distally towards margin; length of free adcau-
line wall, 0.244036 mm. Margin, 0.15-0.18
om in diam, One hydrotheca shows regenera-
tion after breakage at base.
Remarks: The Pearson I, material compares
well with Bale's (1888) specimens from Port
Dennison and Port Phillip Bay. bat they have
HY DROLDS
a shorter length of hydrotheca free of the inter-
node than Ritchie’s (1911) specimens from
Wata Mooli, N.S.W.
This is only the fourth record of this
ipparently rare deeper water species, and a new
record for S. Aust,
Symplectoscyphus sobdichotomes (Kirchen-
pauer, 1884). Ralph, 1961a: 813, fig. 20,
Sertnlarelia suhdichatoma Kirchenpauer, 1884:
46, pl. 16, fig, 1. Bale, 1Y¥I4a: 20 (discussion).
Records: R, 30 m, on bryozoa; Sto. F. 65
m, on bryozoa,
Marevial> Two colonies, each of a few infertile
stems. Siemy straggling, to 4 cm long, alter-
nately, but irregularly branched. Nodes present
on stem and branches, stem internodes indis-
tinct, 1-3 annular constrictions at junction of
hranch with stem: several branches terminating
in tangled anastomoses. Hydroshecae somewhat
conical, with & slight concavity in middle of
abesuline wall; a fine diagonal line running
From the base of the adecauline wall to a small
internal pee in the flexure af the abcauline
wall,
Remarks; The Pearson |, specimens compare
closely with microslides of Sertularefla divari-
cata from the Great Australian Bight in the
collection of the NMV (Bale 1914a. p, 20).
Svmplectoscyphus neglectus (Thompson, 1879).
Shepherd & Watson, 1970; 140,
Sertutlarella neglecta Thompson, 1879: 100, pt,
15, fig. 1. Bale, 1884: 110. pl. 3, fig. 3, pl. 19,
fig. 22, 23; 1915: 287. Blackhurn, 1942: 115,
Syinnlectoseyohus sp. Ralph, 1966: 163, figs.
~4
Records: R, 25-30 m, on Delivea pulehra,
Metaconfolithon charoides, and other red
and hrown algae.
Material: Luxuriant fertile colonies. Stems to
2 om long, beginning with 2-3 oblique proximal
twists, branches suballternate, no secondary
branching, but occasionally a branch produced
into a tendril. Hydrothecae triangulat in see-
finn in young stems, with 3-4 distinet annular
ridges, thecal walls of mature specimens much
thickened and rounded, the annulations less dis-
tinct: marginal teeth of younger hydrothecae
long and sharply pointed, blunt in older speci-
mens. One-3 smal) internal submuarginal teeth,
sometimes not developed, but in mature hydro-
theeae may be thickened and projecting into
centre of cell, Gonorhecae abundant, male and
female on separate stems. Female gonophores
borne thickly on stem and proximal part of
branches; male gonophores bome only on mid-
175
region and distal part of branches, Both sexes
on a short pedicel arising beside a hydrotheca;
hady with 10-15 annulations and 2 hollow.
conical distal processes. Female gonotheca
stout, inflated, widest near middle, blastostyle
spindle-shaped, supporting 4 cluster of ova.
Male gonotheca long, narrow, widest near base,
with a short distal obliquely inclined neck,
blistostyle thin, rod-shaped, becoming indis-
tinet distally.
Remarks: Bale (1884) in his redescription of
8. neglectus, had only dried material before
him, and inferred the transversely wrinkled.
tiangular hydrothecae to be artifacts of drying.
However, much of the present material, par-
ticularly the younger stems, shows this to be
anormal character of the species.
Bale’s surmise that 8, neglects would show
sexual dimorphism is demonstrated hy the
Pearson I, material. His figure “a” (PI. XTX,
fig, 23) with “two latge conical hollow teeth,
one more elevated than the other” ts a female
gonotheea, and gonotheca “h" (fig. 22) with
“teeth smaller and about equally elevated” is
male. Tn the Pearson lt. material, the tooth-like
conical processes of the female gonothecae are
of approximately equal hetght, while those of
the male are of unequal length and fairly short,
hat this is variable througheut the range of
specimens. Ralph (1966) described and figuied
Symplectoveyphus sp. (from Port Phillip
Heads) with smooth hydrothecac of triangular
section and sharply erect marginn) teeth.
Although the hydrothecue are smoother than
usual, the two latter characters clearly distin-
guish her material as young specimens of S.
neplectus,
5S. neplectus is a very common epiphytic
hydroid in southern Australian waters, occurr-
ing on a variety of algae, Ii is easily recusmised
in the ficld by the incurved habit of the
branches, the encfustation of pink coralline
ilgae usually present, and the bright ycllow
gonothecav. The short marginal teeth seen on
alder hydrothecac are probably the result of
constant abrasion against other slems and algae
in the very turbulent conditione in which it is
usually found.
Symplectoscyphus indivisus (Bale, 1882).
Raiph, 1961a; 803, fg. 15. Shepherd &
Watson, 1970: 140.
Sertataretia indivisa Bale, 1882: 24, pl. 12, fig,
7; (8&4 105, pl. 3, fig, 4, pl 19, fig, 27; 1915-
285, Biackburn. 1942: 115.
Records; KR, 21-45 m, on Lauretivia elata
aml Sargessam spinuligerum: S$, 12-30 m,
176
on Sargessan spp, Déistreniiune sp. and
Polvsiphenia sp-
Material: Colonics abundant, 1 colony tertile,
growing from a loosely wound hydrorhiza.
Stems to S mm long; gonothecae clustered
thickly at base of stems. Colour—bright yellow.
Remarks: The colonies fall within the known
range. of variation of S. indivisus, but are
separable into 2 distinet morphological
groups. The first group comprises stems with
short, strongly undulated internodes, often com-
pletely occupied by the hydrotheca, The hydro-
thecae are inflated, irregularly undulated. with
a short submarginal neck, The gonothecae are
squat, deeply crumpled, with very short pedi-
vel, This group compares with Bale’s (1888)
5, fudivisa trom Portland, Vic. and figs. 5. 6, of
§. varigbilis from Port Jackson. N.S.W. The
neck region of the Pearson [.. specimens are
however, more contracted than those figured by
Bale.
The stem internodes of the second group are
longer. and both internodes and hydrothecie
are less inflated. than those of the first group-
The thecal wall is only occasionally faintly uo-
dulated, and the neck region Is longer, The
colonics ure infertile.
There ws thus a goal correlation between
stem morphology and environmental condi-
tions. as the robust fertile) stems were found
only on the rough-water site. whereas the mure
ficxuous (infertile) stems occurred only on the
sheltered side of the island,
Symplectoscyphus pygmaeus? (Bale, 1882}.
Serrvlarctla premaeus Bale, 882: 25. pl, 12,
fig. 4; 1884; LOS, pl. 3, fig. &. pl. 19, fie. 19.
Biackilrn. 1942: 115, Hodgson, 1950: 3h, figs.
63, 64,
Symplectoveyphus pygmaeus (Bale). Ralph,
1961a: £05, fg. 16.
Records: R, 18-30 m, on Herdmania
moms, bryozaa and compound ascidians:
§, 18-24 m. on stem of Thecocarpus divari-
cafes var. eystifera,
Marerial: Infertile colonies comprising a few
stems to 4 mm long. Coleur-—hright yellow,
Remarks: It is difficulr to distinguish between
infertile muteriul of S. pygmaens and &, ren-
roni (Bartlett. 1907), Ralph (1961) distin-
guishes between the two species on the line of
fine dots passing from the base of the adeauline
wall to a point one third the distance up the
abcauline wall in S. pygmaeus, and a lower
diagonal in §. renton?, However. examination
of a series of microslides of 4. pygrnenty in the
collection of the NMY, shows that this is not
JEANETTE E. WATSON
a reliahle distinction, as the height of the
diagonal varies considerably betweem different
stems, and even among hydrotheeae on the
same stem.
Most of the hydrothecae of the Pearson 1.
matenal have # ling of dots joining the ab-
cquline wall about one quarter the distance up
from the hase, a distance vreater than that
given by Ralph as diagnostic for S$. pr prices,
the junction being marked by a thickened noich
on the inside of the wall, As the Pearson I.
specimens most closely resemble 5, pyenraens,
especially one slide in the Bale ¢ollection
(NMV} labelled “Queenscliff, 1881", the speci-
mens are provisionally assigned to this species.
Symplectascyphus macrothecus (Bale, 1882).
Shepherd & Watson, 1970; 140,
Sertulerela mavrofhecn Bole, (882); 25. pl 13.
fig, 1; 1X84; 107, pl 3, fe 4. pl 19. fig. 24.
Bartlett, 1907: 65, fig.
Records: R, 24m, on Acrocarpia paniculate,
Materials One infertile colony of a few stems.
Hydroerkiza «a coarse: undulating tube, Sremi
to 4 mim long, robust, atheeate part very short.
with a strong distal constriction. Jnternades
conspicuously inflated’ behind hydrotheca,
nodes sharply twisted. Aydrothecue large, 2-6
on ster, completely occupying internode; fixed
adcauline wall 0.18-0.20 mm. free adewuline
wall 0.35-0.40 mm; abcauline wall 0.35-0.40
mm. Thecal wall smooth, with a noteh on sh-
cuuline side below margin, opposite the sub-
marginal tooth. Margin 0.15-0.20 mm in diam,
tnieral view, Three internal submarginal teeth,
the aheauline tooth best developed.
Remarks; The present material agrees well with
deseriptions snd figures by Bale of §. macra-
theca.
Symplectoscyphus rostratus 1.5p.
FIGS. 28-35
Type marerial and Records: Holotype, NMV
G1981. microslide--R, 27-30 m, on Surgas-
rum verruculosin, G2095, preserved miat-
eriul, remainder of helatype colony; para-
types, NMV GI982—R, 27-30 m. on Sar-
gauss verruenlusim: GAYSA—-R, 33 m. on
bryozoa; G1984—R, 27-30 m, on Surgayynnt
verruculosunr, mictoshdés: SAM H36——S, 46
m.on ted algac; microslide.
Descriptions from haletype and pérarypes:
iHydrorhiza tubular, loosely adherent to sub-
strate, Stems short, to 2 mm long, unbranched,
bearing I-3 hydrathecve, perisare thick and
brite. Stem internodes twisted, Inflared behind
HYDROIDS "iy
hydrotheca. proximal internode with 2-3 annu-
lations; width of internode at base of hydro-
theca O:.17-0.26 mm. Aydrotfycae alternate,
occupying most of internode, directed towards
front of stem. cach succeeding internode aris-
ing behind base of preeeding hydrotheca,
directed ovtwards, giving stem a zig-zag
appearance, Hydrotheca barrel-shaped, narraw-
ing ta margin, with 2-3 shallow annular ridges
passing completely around mid-region of thecal
wall, Depth of hydrotheea (from hase to mar-
gin) 0.42-0,50 mm; 0,27-0.30 mm. in diam. at
widest part. Margit contracted, small, rim
heavily thickened, with 3 tecth—1 blunt tooth
in the ceatral adcauline position, forming a
raised beak-shaped crest: 2 blunt. tow lateral
tecith, flanking crest. Abcauline side of margin
a shallow curve, Margin 0.12-0.16 mm in
diam. (lateral view). 1.10-0.13 mmr high (ad-
cauline embayment to crest). Aperture facing
obliquely outwards, depressed into the distal
ridge of the abcauline wall. Four internal sub-
Marginal teethe-2 adcauline, long, flanking
Taarginal crest, projecting downwards into cell;
I. low and ledge-ike (not well seen in an-
terior view) just below margin in the centre
of the abcantine embayment, and 1, similar in
shape, but smaller, deep in the adenuline side
of thecal neck. directly opposite the alcauline
submatginal tooth, seen only in lateral view,
Hydraeth with abeauline caecum, connected
by u delicate web to the abcauline wall belaw
the internal tooth. Gonothecae large, ovate, 0.8
mm long, 0.6 mm wide. arising from a short
pedicel hetow proximal hydrotheca, with 5
strong, crumpled annular ridges, and 3 low,
fairly sharp apertural teeth,
Remarks: The material of S. rastrams from
Pearson I. included only 2 gontthecae, one of
which was immature.
S, rostratus is in some Tespects transitional
between the smaller forms of 5. indivisns Bale,
and S$. snacresiecis Bale, resembling the former
in géneral aspect af |he trophosome and gono-
some, and the latter in the arrancement oF the
imMerrial submarginal teeth, There ts hawever,
only | abcauline submarginal tooth in S, ree
fratus, compared to 3 in J, macrotherus. fn ¥.
indivisns, the 3 mareinal teeth are alternate
With the tnaruinal teeth, The raised adcautine
crest further distinguishes §. resiralys.
Symplectuscyphus epiznicus n.sp.
FIGS. 31-33
Type Material and Records; Holotype, NMY
G1985, microsiide—S. 20 m, on Theco-
rarpus divaricanns van cvatifera; G2096, pre-
served material, remainder of holotype co-
lony; paratypes, G1986, GI9R7T, GLOSs,
microstides—R, 30 m, on TL divaricarer var.
evstifera; SAM H33, micrastide,
Description fram heletype and peratypest
Hydrorhkiza tubular, of same diam. as sterns.
Stems simple, short, unbranched, to 7 mm long.
Proximal stem mternnde short, athecate. with
4-5 annulations, width 0.13 mm; following
internodes of variable Jength. 0.50-1.05 iim,
ftydrorhecaé large, pecisarc delicate, alternate.
in one plane, a maximum of 4 on stem. nat
immersed in internude, distal on long inter-
codes, occupying almost the whole Jength of
short internodes, without definite floor. barrel-
shaped, widest about middle, narrowing only
slightly to margin. Leneth of fixed adcauline
wall 0.22-0.28 mm, free adcauline wall 0,21-
0,30 mm, ubcauline wall 0.45-0.53 mm. Mar-
gin O,28-0.31 mm in diam., thickened, with 3
equi-distant bluntly pointed tecth—1 edceuline,
2 lateral abeauline. Operculum of 3 delicate
flaps, No internal submarginal teeth. Hydranth
with approx. 24 tentucles, and an abcauline
cuecum, Gonetiecae large, ovale, neatly 3
times length of hydrotheca, 1.11-1.35 mm
long, widest at top. 0.87-1.02 mm, tapering to
a short pedicel arising below proximal hydro-
theca, walls fuintly undulated proximally. the
anmulations more distinet about mid-region.
Aperture small, circular, 0.13-0.15 mm in
tlam., depressed into the most distal annula-
tion of the gonéthecal wall; 4 very low rounded
teeth, Gonophores femule, mature, now filling
Pponothecal cavity, with 10-16 eggs, Colanim—
yellow.
Remarks; The colonies arise from a single sta-
fon running up the main stem and branches of
the host, the stems and occasional single hydro-
thecae given off at irregular intervals.
S. eplzeicus resembles one of the farger
forms of §,. indivisns (Bale) (ie. “Sernularella
feriahiliy’ Bale, 1838) in size and structure,
and could easily be confused with this species
in preserved material, However, S. epizotcus
lacks the internal submarginal teeth which dis-
tinguishes 5. indivisns.
Sertularia meacrocarpa Bale, 1884: &0, pl. 5,
fig. 2, pl. 19, fig. 11: 19l4a! 14; I19[5:
277, Mulder & Trebileock, 1914b; 42,
Hodgson, 1950; 27, fig. 47. Shepherd &
Watson, 1971: 140,
Records: R, 25-34 wy S, 4-25, anvone hol-
fasts of red algae.
178 TRANETTR F, WATSON
Material: Colonies abundant, growing in thick
tangled clusters. Stems us 12 cm long, infertile
except for | gonotheca. Colerr—dark hrawn.
Remarks: This distinctive species is easily
recognized by its dark colour, and large.
tangled! colonies which usually grow at the base
of ulgue in semi-sheltered situations.
Serfularia unguiculata Busk, 1852: 394. Bale,
L884: 76, pl. 6, figs. 9-12; L894: 100;
L9]4a: 16; 1915: 273. Blackburn, 1942:
113. Hodgson, 1950: 26, figs. 45. 46.
Ralph, 1961la: 788, fig. 13,
Records: 8, 25 m, on Herdmanta momus,
and among algal holdfasts.
Material; Several large infertile colonies. Stems
short. to 3.5 cm, unbranched, Proximal branch
internodes with 3 pairs of hydrotheeae, euch
succeeding internode with 2 pairs of hydrothe-
cae, nodes. indistinct. Ayedrothecae on branches
adnate for two-thirds of length; cauline hydro-
thecac not immersed in stem. Colour—orange
brown,
Remarks; Although shorter, the stems m the
present collection conform to Bale’s (1884)
description of the thick stemmed, long Inter-
node form of $, uaguienlate.
Although relatively common at Pearson L, 9.
wunrguiculata was restricted to sheltered water.
Sertnlaria bicuspidata Lamarck, 1816: 21.
Blackburn, 1937: 367.
Serthlaria bicornis Bale, 1882: 22, pl. 2, fig.
3) 1884: 83, pl. 5, fig. 9.
Records; RB, 45 m, on Rhodopeltis australis
und Metamastophora flabellata,
Material; A few infertile stems. Stems sttilty
erect, branched, to 1 cm long, Colour—dark
brown,
Remarks: The distinctive paired finger-like pro-
cesses flanking the margin distinguish this spe-
cies From all other species of Sertularia in Aus-
tralian waters,
This is the first undoubted record of §. his
cuspidara from S. Aust, (Other localities—
Queenscliff, and Lady Julia Perey 1., Vic.)
Sertularia maccallumi Bartlett, 1907: 62, fg,
Mulder & Trebileock. 19144: 7, pl. 1, figs,
3. Bale, 1919; 340. pl. 16. figs. 3, 4,
Shepherd & Watson, 1970: 143,
Records: R, 25-45 m, on Carpopeltic pfiyl-
lopora and Prerocladia licédet.
Muterlaly Luxuriam fertile colonies thickly
oyer-running algae. Stems to 4 mm long.
Sertularia acuta (Stechow,
Colowr—hydrorhiza brown, hydrocaulus bright
yellow,
Remarks: §. maccallumi 15 one of the com-
monest. hydroids in the Pearson J. collection.
Although both species of algae were also
recorded at West L. (Shepherd & Watson 1970),
S. wraccallumi was never Found on P, lucida,
and only occasionally on C. phyllophera at that
locality,
1921), Millard,
1958; 192, fig. 8, Shepherd & Watson,
1970; 140.
Sertularia loculosa Bale, 1884: 9), pl, 4. figs.
5,6; (913: 121, pl. 12, figs. 7, 8; M15: 272.
Tridentata acuta Stechow, 1921; 231.
Sertularia balet Briggs, 1922: 150,
Records; R, 45 my, 8.15 m, on Srenecledia
australis.
Marerial; Abundant fertile colonies, Stems un-
branched, fo 5 mm long, internodes with an
oblique proximal, and a transverse distal joint;
exceplionally, a transverse joint is followed by
2 oblique jomts, Celeyr—yellow. Gonorhecae
with 4-5 deep annulations,
Reinarks: The Pearson 1, specimens correspond
closcly to the short-celled form of “S, loculosa"
Bale.
Millard (1958, p, 198) distinguishes S, acuta
from S, turbinata (Lamouroux, 1816) in her
South African material partly by the presence
of transverse stem nodes in the former species,
and oblique nodes in §. rurpinara. The Pearson
T, material has both transverse and oblique
joints on the one stem, thus further reducing
the difference between these two closely related
species. 8. acuta is a common epiphytic species
in the collection and is associated with only one
specics of alga.
Amphisbetia maplestonei (Bale, 1884), Rees &
Thursficld, 1965; 142. Shepherd & Wat-
son, 1970: 140.
Sertularia maplestone’ Bale,
fie 4, pL 19, fig. 2
Serfularia bidens Bale, |884:70, pL 6. fiz.
19, fiz. 1; 1914a: 14,
Records: R. 34 m; 8, 25 m, among
holdfasts.
Material: Luxuriant fertile colonies.
flexuous, 5-12 em tong.
L884: 70, ph 6,
6, pl.
algal
Stems
Remarks: The distal conical processes of the
gonothecae are modetately well developed, bul
many gonothecae have only 1 process on the
abcauline side.
TY DROIDS (74
Amphishetia pulchella (Thoinpson, 1879).
Shepherd & Watson, 1970: 140.
Serndana palehelia Thompson, 1879: 109, pl
TX, fies, 3, 3a, Bale, 1884: 71, pl. 6, fig. S, pl
19. tig. 10.
Sertularia
1942; 113,
Records; R, 45 m, on Praracladia lucida; §,
14 m, on bryozou epiphytic on algac.
Marerial: A few infertile stems to | cm long,
Remarks: Blackburn (1942) included A. pui-
ehella in the synonymy of A, maplestone? al-
though A. pulchella is a prior numeé. Although
similar in microscopic details of the tropho-
seme, the lwo species seem to be distinct, und
are readily distinguished by the gonothecue and
size of the stem. The stems of A. nwplestere’
ure long and robust, while those of A. pulehella
rarely exceed 2 cm. Both species are epiphytic;
the larger species, A. smaplesronei, is confined
to the hasal parts of algae, while the more deli-
cute A. pulehella epiphytises the fronds. Pos-
sibly, further work may prove the two tu be
ecomarphs of the one apecies.
maplesioned (Bale), Blackburn,
Amphisbetia olseni nsp.
FIGS. 34-37
Type Material and Records; Holotype, NMV
G2001, microslide—R, 33 m, on sponge;
G2097, preserved material, remainder of
holotype colony; paratypes, G2002, G2003,
32004, microslides—R, 33 m. on Herd-
munta moms: G2O0S—S, 17-33 m, on
brown algae, microslide: G2098, temainder
of paratype colony G2003, G2006-—R, 33
m, on red algae, microslide: SAM H34-—R,
33 my, on Sponge. microslide.
Description fram holotype and paratypes:
Hydrorhiza tubular, Stems to 7 mm long,
monosiphonic, stiffly erect, branched. Proximal
stem internodes athecate, terminating in a
strong V-shaped joint. suceecding internodes
thecate, nodes indistinct, but if present. V-
shaped, slender. Internodes 0.52 mm Jong,
diam. at node, 0,06-0,09 mm; 3 hydrothecie
on stem internodes, 1 axillar. 2 subopposite,
Branching regularly alternate, up to S branches
on stem, atising from a long proximal apo-
physis given off at 70° to stem. First branch
internode very short, athecate, with a transverse
proximal, and V-shaped distal joint; remainder
of branch without internodes, but with wp to
5 pairs of hydrothecae. Hydrothecae on
hranches subopposite. in 1 plane, adnate
approx. two-thirds of their lenzth, saccate,
widest near middle. narrowing 2 margin, Fixed
adeauline wall 0.12-0.16 mm long, free nd-
cauline wall 0.05-0.08 mm, standiag our hori.
vontully, of at a slight upward angle from the
internode, Abcauline wall 0,16-0,19 mm, a
pronounced inflextire about one-third the dis-
tance up from the base, followed by 4 sharp
outward bend, but this may be reduced te a
mere concavity in the abcauline wall, Paired
hydrothecae on younger parts of stem and
branches in contact along fixed adcauline wall,
but separated in older parts of stem) if in con-
tuner, the adcauline wall is straight, otherwise
it is bent parallel to the inilexure of the ab-
eauline wall. Margin with 2 long sharply
pointed lateral teeth 0.07 mm long, separated
by a deep, almost horizontal abcauline sinus;
adcauline wall indented behind matgin. Oper-
culum of 2 flaps, abcaulinc component fixed.
Aydranth with approx. 12 tentacles, Gono-
thecae arising from lower stem behind proximal
pair of hydrothecae, large. ovate, fattened,
1,02-1,32 mm tong. Perisare thick, walls
smooth, widening distally to a shoulder 0,84-
1.02 mm width, produced inte a pair of short,
almost laterally directed spines. Aperture citcu-
lu’, 0.35 mm. in diam., with a slightly raised
collar and a ring of minute internal denticles.
Operculum a. circular flap. Gonothecae empty.
Colour—light straw colour,
Remarks; The branching in 4. olseni is regu
larly alternate, where a branch fails to develop,
the stem internode is longer, and has I axillar,
and 2 pairs uf subopposite hydrothecac, then
branching resumes again, The branches are
very brittle and break off eusily wt the slender
proximal joint.
A. olseni 1s closely reluted to both A. prl-
chella (Thompson) and 4, tidens (Bale) in
structure of the trophosume and gonosome, but
is distinguished from these species by the
straighter, Jess flexuous stem, the pronounced
concavity of the abcauline thecal wall, the less
prominent marginal teeth, the greater propor-
tion of the hydrothees adnate to the internode.
and the shape of the gonotheca,
With one exception (8, 17-33 m), the colo-
nies were all found within the same area on the
rough water site.
A_ olseni is named after Mr. A, M, Olsen,
whose interest and encouragement has done
much to foster marine science in South Aus-
tralia.
Amphisbetia minima (Thompson, 1879),
Ralph, !96far 774, fig. & Shepherd &
Wilson, 1970; 140.
FIGS. 37, 38
180
JEANETTE E. WATSON
Figs.
Figs,
Fig.
Fig.
Fig,
Fig.
wu SO
wus"
38 39 40 4|
31-33. Symplectoscyphus epizoicus n.sp. Fig. 31—Whole stem with gonotheca. Fig. 32 Hy-
drotheca enlarged, anterior view, with hydranth. Fig. 33.——-Hydrotheca, lateral view.
Drawn from holotype,
34-37. Amphisbetia olseni n.sp. Fig. 34.—Whole stem, from holotype. Fig. 35, part of stem and
38.
39,
40,
41,
branch, enlarged. Fig. 36,—Part of stem and gonotheca, from paratype. Fig. 37.—
Hydrotheca, enlarged,
Amphisbetia minima var. pumiloides Bale. Part of stem with two internodes.
Amphisbetia minima var. intermedia Bale. Part of stem with two internodes, Figs. 38
and 39 drawn at same scale for comparison.
Amphishelia minuscula Bale. “Short internode form".
Amphisbetia minuscula Bale. “Long internode” form.
HYDROIDS tat
Sertularia minima Thampson, 1879: 104, pl 17,
fig, 3, Bale, 1882; 21, 45, pl. 12, fig. 25 1884:
$9, pl. 4, figs. 9. 10, pl. 19, figs. 12, 13; 1945;
269; 1924: 248. Mulder & Trefileack, 1914b:
39, Sreehow, 1925. 231, fp. K. Blackburn,
1942: L14. Hadgson, 1950: 23, figs. 41, 42,
Millard, 1957; 221.
Records; KR, 12-50 m;S, 15-25 m, on algae
{see Remarks),
Material) Luxunynt fertile colonies, Stents to
35 mm long. Colow7e—tight brown.
Remarks; Bale recognized 3 varieties of A-
wine jn southern Australian walters, the
largest, var, pumtloides Bale, 1884 (from
Queenscliff, Vic.), a “typical” farm Irom Port
Phillip Bay and New Zealand waters, aud var.
iermtedia Bale, 1915, from the Nuyts Acchi-
pelago in the Great Australian Bicht. Ralph's
(1951) figures und description show her New
Zealand specimens. to be closely allied to the
Australian typical form, but there are nemato-
thecac scattered throughout the hydrocautus, a
condition not normally encountered in the Aus-
Uulian material. 4 minime is one ol the most
abundant hydroids in the Pearson I. collection,
und the material exammed falls with little inter-
vradation into 2 of the varieties, var. pudai-
loides und var. intermedia, Although difficult
to distinguish in preserved material, the vyarie-
hes are casily separated in mounted prepara-
tions. These varieties have not previodsly been
recorded together in the one Jocality.
The present material corresponding to var.
puniufoides is a robust form, conforming to
Bale’s (1884) ceseription. The bydrothecye ure
almost entircly adnate, and the typical wedge
af perisurc belween hydrotheca and internade
is well developed. The gonothecue are variable
in shape, round Lo clongute mm Jatetal view;
those with # raised apertural collar do not have
wring of internal submarginal denticles. The
var. intermedia, not figured hy Bale. as much
smaller, with shorter internodes, and rectilinear
hydrethecae. Nematothecae, varying from
short cylindrical tubules to mere breaks in the
perisarc. are present on all stems, but are cnn-
fned to the infrathecal chamber of the proxi-
faal hydrothecae. Gonothecae sure round [0
ovate, Dimensions of the 2 varieties from
Pearson !. are given for comparison.
var. var.
_ Dimensians. mat pamileides __fatermecia
Internode length 1.35-0,38 A.25-0,37
Digan. al node 0.05-€.08 0.03-0,04
Livdralhecu Tength 24-029 .t7-1.22
With few exceptions (Rilchie I91T), A.
mninwe is recorded as an epiphytic species, and
at Pearson [, the 2 varieties show a stroug
selectivily towards certain species of algae. The
var. puumiloides was found only on the robust
brown aleae Sargassum bracteolosum, 8,
varias, and SY. verrucu/osum, as well as Aero-
carpla puniculeta; var. intermedia Was asso-
ciated with the more delicate red algae Rhoddy-
tnenia australis, Metamasiophera flabellata,
Laurencia elata, and Carpopeltiy phytlophora,
with one record on the delicate brown alga,
Distromium flabellatum. Shepherd & Watson
(1970) noted that many commonly epiphytic
hydroids show varying degrees olf preference
for particular species of algal substrate, but a
differential selectivity by varielies of the same
species has not previously been recognized.
Almost all the colonies have the pegged
hydrorhiza typical of A. minima. Bale (1915)
and Mulder & Trebileock (19I4u) noted this
tact; Ritchie (1931) suggested it muy be a
response 10 Wave action, but Ralph (1961)
could find “no constant relationship vo enyiron-
mental conditions” to accvunt for the thicken-
ings, In many of the colonies of both varieties
of 4. ntinima from Pearson L, the hydrovhtes
is often tubular and loosely winding when in
comuct with the curved and cylindrical sur-
faces of the lower stems of the ulga, then flat-
tening out and developing the transverse murk-
ings as the stolon passes onto (he broader
fronds. This change in cross section of the
stolon may {herefore be etther a response to the
greater movement of the algal frond in turbu-
lent walter, or it may be related to the narure
of the algal surface.
The systematic status of the 3 so-called varie-
ties of A. minima, and of the whole “A.
minha? group, including 4. avniscala Bale,
A. fureaie Trask, and A. stuellevi? Bale, needs
further clucidation. Tt is possible that all may
be ecologic variants of the one species, or
several distinct. bur closely reluted species.
Amphisbetia minuscula (Bale, 1919),
FIGS, 40, 41
Sertifarla minima var. tebathere Mulder &
Trebileuck, 1934b: 40, pl. 4. fig 1-
sSeesatarte puisifla Bale, 1915: 271, pl. 46, figs.
i=
Sertifaria minuscila Bale,
burn, 1942; 114,
Records: R, 30-40 m, on Lauvrencia elite,
Disttomiun flabellaiuin, aod Herdmeania
moms: S, 25-31) m.ion the stem of Cauuvlerpa
sp.. and Halicernaria longirostriy.
Morera; Abiadant colonies, some fertile,
Stemy simple, to 5 min) long, internodes vari-
able. The slems ace divisible into 2 groups—ag
1919; 340, Bliack-
182
long internode group, internodes 1),42—0.44 mm
long, anu « short internode group, internodes
0.30-0,.34 mm long. Nodes of both stem groups
0.06 nim wide, indistinct, transverse, occa-
sionally a V-shaped joint in distal region of
stem. Hvedrethecae similar to deseriptiens al
authors, fixed adcauline wall 0.19-0,22 mm,
free adcauline wall G.07-0,11 mm; abcauline
wall 0.19-().26 mm; an indistinct downwardly
curved ridge passing back from the embayment
between the marginal teeth into the junction of
the adcauline wall with the internode. Gono-
thecae 1.20-1.26 mm long, excluding pedicel,
maximum width, 0.75-0.97 mm, present only
on the “short internode” form.
Remarks; The hydrothecae of the “long inter-
node” form of the Pearson I. material are larger
than those of the “shart mternode” form, but
both ate smaller thun measurements from
microslides of 4. miinuscula, and the type of
“4 minima var. tubathece” in the collection of
the NMY. The “long internode” form conforms
with measurements ot A. pusilla (Bale), while
the “short internode” form is similar to the
var. tubatheca of Mulder & Trebilcock. ‘The
nemitothecue noted by Bale and Mulder &
Trebileock are present in only a few of the
proximal sium internodes of the Pearson I.
specimens; the intrathecal ridge, noted it the
present material, is not present in the type. A.
nunascula displays a wide choice of substrate,
Some correlation evidently exisis between stem
type and environmental conditions, since the
“long. internode” form was abundant on the
sheltered site, whereas the more robust “short
internode” form was found only on the rough-
water side of the island. This suggests that deve-
lopment of a thickened stem with short intet-
nodes is advantageous to withstand rough water
conditions. :
Family PLUMULARILDAE
Pycnotheca producta (Bale, 1882),
Plumularia praducta Bale, |882; 34%, pl. TS,
fig. 3; 1884: 133, pl, (0, fig. 4; 1894; FTI.
Kirehenpuueria prodicta Bale, 1914a; 59, 1915;
302. Blackburn, 1942: 107.
Records: R, 24 m. on Distromitin flubetla-
wm; 8, 24 m, on Avmenera?,
Material A few scuttered infertile colunies.
Stemy to F mim lovg.
Remarks: The material contorms exactly tu
Bale’s description of P. prodnete.
Antennecila tubulosa (Bale, | 894),
FIG. 42
Piganiasia tihalesa Bale,
hes. 2-5,
1894: 114, pl 5,
JEANETTE FE. WATSON
Records: R, 27-30 m, on bryozoa: S, 26-30)
m, On Sargassum sp.
Material: Seyeral colonics, cach comprising a
tew infertile stems, Stems to 3 mm lone, ansing
from a thick hydrorhiza. First internode with
a proximal constriction, fllowed by an athe-
cate Internode with 1-2 nematothecac and an
oblique distal node. Athecate internodes 0,1 1—
0.16 mm long; thecate internodes as described
by Bale, 0.27-0.37 mm Jony, with 1 median
nematotheca and 2 scoop-shaped lateral nenia-
tothecae, Aydrothecae long, 5-G on a stem,
proximal part tubular, perisarc thick, adcauline
wall 0.25-0.27 mm long; abcauline wall 6,22—
0.30 mm long. distal part of adcauline wall
more convex than abcauline side, Margin 0,13-
0.15 mm in diam., deeply sinuated, curving up
to meet produced adcauline wall. Co/our—
yellow.
Remarks: The Pearson L maternal, althougl
definitely reterable to 4, rubrilose, nevertheless
shows considerable variabilicy in thickness of
perisarc, length, diam, af inlernove, and shape
of hydrothecae. Bale (1894) comsidered that
ef, tebdlosa may be a variant of A. canipannla,
or jhe “Antennella” form of an unknown Plu-
mularian,
As the present matenal shows no sitn of
branching, 4. tubulosa may therefore be con-
ssdered 4 distinct spectes.
This is the second record of 4. rebudosa, anc
a first record for S, Aust. (Other locality—Pon
Phillip Bay, Vic.)
Antennella campanuliformis (Mulder & Trebit-
cock, 1909),
FIGS. 43, 44
Pienvlocia campanatiformly Mulder & Trebil-
cock, 1909F FL, pl. T, figs. 6, 9, 10; T9LT: 115.
Records; R. 30-45 m, on Sargassien sp.
Lenvrencta elate and Preroelacie lucida.
Material: Colonies common, fertile, The spéci-
mens contorm to the type microslide of Pfi-
mularia campanuliformis in the collection of
the NMY. The following description supple-
ments that of Mulder & Trebileock,
Hyvdrorhiza tabular. Stems to. 1 cm long, first
stem internode 0.5 mm long, athevale, with 1-2
hematothecae and 1-2 proximal annulations,
distal node oblique, following internodes qlter-
nately thecate and athecate. Thecate internodes
0.39-0.45 m Jong, almost entirely oceupied by
the hydrotheca, with an oblique proximal node,
and a transverse distal node, 0.06-0.09 mm
wide. often indistinct; athecate internode short,
O.2R8-0.31 mm (measured along base of hydra-
HYDROIDS 183
cladium). Hydrarhecae cup-shaped, deep, base
curved, abcauline wall 0.20-0.22 mm tong,
concave: adcauline wall 0.21-41.25 mm long,
straight or wath » slight convexity below the
margin, both walls thickened, the abcauline
flange eXtending back tu the median nemato-
theca, Margin circular, 0.21-0.23 mm in diam.,
slightly »sinuated, Nemetorhecae large, bithala-
mic, O.08-0.L0 mm Jong, distal cup 0.05 mm
in diam, 3 on the athecate internode—]
median, base stout, cup excavated on adcauline
side. 2 laterals below bydrotheca on a very
shart apophysis of the hydrocladium, cups nur-
row. txcuvated on inner side, sides slightly in-
rolled; | on athecate internode, similar ta
other's; 2 similar in shape. but larger than
cuuline nematothecae on the pedicel of Female
gonotheca, facing outwards, cups excavated on
the side facing gonotheca; 1 nematotheca on
male gonotheca above pedicel. Gonothetae of
both sexes on the one stem, arising beside the
median nematotheca on theeute internodes.
male small, 0.22-0.27 mm long, ovate. 0,10-
0.14 mm wide, only on proximal stem inter-
nodes; female large, 0,64-0,66 mm in diam.
globular, only on distal internodes; female
gonophore of | large egg surrounded by a top-
shaped blastostyle.
Remarks: Although Mulder & Trehileock des-
eribe and figure the hydrotheca of 4. campo-
nutiformis as campanulate, this is somewhat
misleading, as the type specimens as well as the
Pearson [, material have almost tubular hydro-
thecae.
This is the second record of A. canrpmiuli-
fermis, and a first record for S. Aust. (Other
locality—Vic_}
Antennella secundaria (Gmelin) s. sp. dubia-
formis (Mulder & Trehbiloock, 1910),
FIGS. 45, 46
Plumelaria debiaformis Mulder & Trebiloock,
1910: 119, pl. 2. fig, 7.
Anrennelfa secuytidarin (Gnielin). Billard, 1913:
8.
Plamularia stcundarja (Gmelin), Blackburn,
938; 346),
Sehizetriche secundaria (Gmelin), Blackburn,
1942: 108.
Records: §, 17-27 m, on compound ascidian.
Rhodymenia austrulis, Distramiam flabella-
tam, Sertuluria unguiculara and sponge.
Material; Abundant fertile colonies. Mydrarhiza
tubular. Sterns lo 6 mm Tong, perisare delicate.
First stem internode Jong, athecate, with 3
nematothecae and oblique distal joint. inter
nade oceasionally with | branch, Following
internodes allernately theeate amd wthecate,
athecate internodes 0.3 mm Tong, with an in-
distinct transverse and a strong oblique distal
joint: thecate internodes shghtly longer, 0.30—
0.35 mm long. Hydrothecae campanulate,
0,20.-0.22 mm deep, set an un angle of 45° to
hydrecladial axis, base flat, abcauline wall
slightly thickened, udeyuline wall almost
entirely adnate, free part closcly adpressed to
internode. Avarefm entire, delicate, 0,26-(,30
mim in diam. Nematothecae as described for 4.
secundaria, 2 present on uthecate internode.
One small suprathecal nematotheca usually, but
not always present in the sinus above the hydro-
theca. Genothecae—male and female on the
same stem, arising beside the median subhydro-
thecal nematotheca, tapering (o a short pedicel.
Female globular, flattened, 0.52-0 58 mm long,
widest. near niiddle. 0.40-0.44 mm maximum
width. cloycd by a thin operculum: 3 nemato-
thecne similar to laterals, in hasal region. Male
gonotheea small, 0.15-0.20 mm long, 0.12-
0.13 mm wide, with 1 proximal nematotheca,
Remarks: The present specimens are identical
with 2 microslides of [tagmentary infertile ma-
teal of Plamularia dubiaformis Mulder &
Trebilcock if whe collection of the NMV.
Mulder & Trebilcock. because of poor material,
were unuble to establish the presence or ab-
sence of the supratheeal nematothecae in P.
dudiaformiy. These nematothecac are clearly
visible in the Pearson [. material-
T have also compared the present material
with fertile material of A. secundaria trom
Mossel Bay; South Africa. provided by Dr
N. A. H. Millard, and with (he exception of the
2 median nematothecac on the athecate inter.
node in the Pearson |, specimens (f in the
South African material). the twa are indistin-
guishuble. As the number of nematothecae on
the athccate internode is not a reliable specific
clitetion, 1 agree with Billard (1913) and
Blackbur (1938) who suggested PL diubia-
fermiy would prove to be a synonym of A.
secundaria.
As the Known South Australian material of
A, secundaria always has 2 nematothecas on
the athecate internode, compared to 1 in the
typical form, und has now been recorded From
two widely separated localities (Pcarson I and
Vic), it Is here recognived as a subspecies of
A. secundaria,
Halopteris suteata (Lamarck, 1816)
Plurnularia saleata Lamarck, (818: 128. Briggs,
1915: 306, pl, 11, fig, 1. Bale. 1914b> 172. pl.
35, figs. 6, 7; 1915: 296.
Plumuteria agliaphenoldes Baie, 1884: 126, pl,
10, fiz. 6.
1x4 JEANETTE
Reeordy; Ro 30m, on sandy floor of cavern.
Material: Several large Fertile colonies. Sten
fuscicled, branched, co 20 em high, growing
from a small fibrous rootstock. Colour-—dtatk
brown.
Reimarks: This species is cusily recognizable thy
its large, erect woody stem, colour, and brittle
texture. At Pearson 1., A. syleata was found
only on the floors of caverns shellored from
gtirse: tL hus however also been noted in open
water to the east, in Tnvestigator Strait (J.W.,
unpublished) at depths of 40 m,
Hulopteris campanula var. campanula (Busk,
1852). Ralph, 1961 6;47,
Plurnelaria carmponita Busk, 1852; 401. Bale,
(884: 124. pl. 10. fig. 5: ISEB: 776; 19t3: 134;
(475; 245, Hodgson, 1950: 40. fiz. 49.
Schizoricha canmpannla (Busk), Blackburn,
1942: 107,
Reeords: KR, 35 m, epilithic,
Murerial; One infertile calony growing from a
common rootstock, Stems —polysiphonic,
branched, to 4,5 ¢m high; some secondary
branching. C'olouwr—yellow.
Renunrks: The specimens agree with descrip-
lions of # canzpanule, and A. campanula vat.
contpunita oF Ralph (1961b)..
Halopteris buski (Bale, (884),
Plamularia bushi Bale, 18845 125, pl. (0, fia.
3. pl. U4. figs, 34. 35; 19)4dur 28, 1915: 296.
Arives, 1915: 304. Hodgson, 1950: 45, fix. 75.
Sehizetricha buski (Bale), Blackburn, 942:
107,
Recardy' RR, 30-338m, on Herdinania
monies, bryozoa and sponge: Stn. F. 65 m.
on werm tube,
Mareriul: Scattered voloniex, euch of iu few
stems ta 2 em long. One stem with immature
inale gonophores
Remarks: The Pearson 1 specimens do not
aliffer significantly from descriptions of [Bile
and Briggs. With one exception (Sin. F) the
colonies are all from the exposed side of ihe
island. They are short and robust. with deeply
incised stem joints; frequently an extra oblique
septal Internodal ridge is developed just below
the adcauline hydrothecal wall, The stems from
less turbulent deeper water (Stn. F) are more
fiexuous, wilh indistinct cauline nodes, and alsu
lack the oblique hydrocladial septa present in
the shallower water specimens. The deeper
water stems were scarlet in colour, Whereas the
shallower water specimens varied from orange
to yellow,
E. WATSON
Halopteris vupposita (Mulder & Trebileock,
1911).
FIG. 47
Plimtlatia appesita Milder & Trebilenck-
1911; 120, pl 2, fig. 5,
Thevoedilus oppositus (M. & T,). Blackbilin,
1938s 316, fig, 2) M42; 17.
Recardy: S, 30 m, on Sargassum spinuli-
gerne and 8. verruculostini.
Mererial: Scattered infertile stems to 12 mm
long, The specimens compare with the type
microslide of Mf. eppeyifa in the collection of
the NMV, and allow a fuller description to sup-
plement the previous brief description of
Mulder & Trebilcock,
Stemy with 2-3 indistinct proximal annula-
tions, followed by a long athecate internode
with 2-3 nematothecae, then alternate thecate
and vathecate internodes of approx, the sarne
length, 0.27-0.39 mm, athecate internades with
a proximal transverse joint and strongly oblique
distal hinge-joint, and 2-3 nematothecuc, if 2,
they ate ane shove the other: the third, if
present, is beside the distal nemalotheci.
Hydrocladia opposite. arising behind the
cauline hydrothecac in middie of the anter-
node, hydrocladia beginning with 2 short athe-
cate internodes, the firs. internode the shorter,
0.95=0,.07 mm long, the second 0.07-0.98 mm
long, with a transverse proximal, and an
oblique distal joint, and occasionally, | nema-
totheca. Thecate hydrocladial internades sinii-
lar ta stem internode. Hydrothecae campinu-
late. 2-3 on hydrocladium, 0.15-0.18 mm
deep, set at 45° to hydrocladial axis. Murgin
0.19-0,21 mm diam., slightly sinuated, rim
eVveried, with a peak on the abcauline side. Ab-
gauline wall 0,12-0,15 mm long, In some
hydrothecue a sinall transverse fold near hase.
Nematothecae 0,05-0,07 mm long, 2 supra-
thecal un stem internodes, 4 on thecate hydro-
eladial internodes—1l median, stour at base.
distal cup cut away on adcauline side, closels
adpressed to internode; 2 lateral stiprathecal.
with slender pedicels, an an apaphysis af tke
internode, extending above hydrotheca, and 1?
small nemalotheca between laterals at hase of
hydrotheea. Hydranth with 10-16 tentacles,
connected to internode by a small orifice in the
upeurve of the abeauline wall,
Remarky: The stem of the type is thick and
robust, and the athecate siem intemodes ave
cunsiderably shorter than the thecate inter-
nodes, with deeply constricted nades-
The Pearson T, specimens have slender stems,
and internodes of neurly equal length, Black-
42,
47.
. 48-52.
HYDROIDS
0.25mm
WU O
5]
O-lmm
Antennella tubulosa (Bale). Part of stem.
» 43,44. Antennella campanuliformis (Mulder & Trebilcock). Fig. 43.—Part of stem with male
s. 45, 46,
gonophores, Fig. 44——Female gonophore-
Antennella secundaria s.sp. dubiaformis (Mulder & Trebilcock). Fig. 45.—Part of stem
with two hydrothecae, Fig. 46—Empty female gonotheca.
Halopteris opposita (Mulder & Trebilcock). Hydrocladium with one hydrotheca.
Gattya trebileocki n.sp. From holotype. Fig. 48—Whole stem. Fig, 49.—Hydrocladium
with three hydrothecae. Fig. 50.—Hydrotheca, anterior view. Fig. 51.—Twin lateral
nematothecae from distal end of hydrocladium, enlarged. Fig, 52,.—Median nematotheca
on thecate internode, enlarged.
sf
hum (1938) also noted this feature in his mute-
ful Erom the Sit Josenh Bunks Group, and pon.
sidered that it may “constitute a distinct
variety”. As the type microslide consists of a
single sten) fragment 3 mm long, il scems pos-
sible that the Sevth Australian maternal nay
better represent The species than the type it-
self.
This is whe third record of Ff. apposita.
(Other localities—central Vic. Sir Joseph
Banks Group, S. Aust.)
Gattya aghovheniaformis (Mulder & Trebil-
cock, 1909),
Plimularia dglaopheniaformis Mulder & ‘Trebil-
cock, 1909: 32, pl. 1, fig. 7
Hafonteris agliapheniuformis (M, & T.), Shep-
herd & Watson, 1970: 140,
Records: R, 18-33, on Phurtuiuria procum-
bens and Callophiyllix coecined,
Muterial: Several colonies of a few infertile
stems each. Mydrorhiza tubular. Scenis to'7 mm
long, bevinning with an athecate internode with
1~3 pairs of couline nematothecae: thecate
internodes. with a proximal hydrotheca, and 2
pairs of suprathecal nematothecac. Hydroctadia
arising from a small apophysis behind hydro-
thee», the first pair opposite, first 2 hydrocladial
internodes slender, short, athecate, 0.04—-0,06
mm long, with transverse joints. distal internode
longer. 0.07-0.09 mm, with 1 medinn nemato-
theea. Mydratheeae 0.17-0.20 mm deep (fateral
view), with broadly lobed margin 0.13-0,18
mm in diam, the anterior and posterior lateral
projections curving inwards Over the aperture,
Remarks: The marginal projections of the
Pearson [, specimens. differ from those of P.
ughophentaformis fiznured hy Mulder & Trebil-
cock. Furthermore, the cauline internodes are
vuriable in Jength, some heing barely long
enough to accommodate the hydrotheca; those
Stems with longer internodes also have a thin
perisarc, are more Hexwous, and have a less
deeply lohed hydrothceal margin Uhan the type.
Following Millard (1962, p. 270), this and
nther species with a toothed thecal margin are
referred to Gatrya,
Gattya balei (Bortlett, 1907).
Plamularia bealei Bartlett, 1907+ 65, Mulder &
Trebileack, 1909; 29, pl. 1, fizs. I-39. Hale:
1919: 344, pl. 17, fig. 6.
Records; R, 14-45 mt on Metamastophara
flabellata, Pterocladia lucida and Ptere-
siphonia?.
Afatertal: Colonies conimon. A few fertile stems
with female gonophores, Sténes te 7 mm long.
JEANETTE E. WATSON
Cojaur—vellow,
Remarks: This rare but distinctive species has
not been recorded previously in S. Aust. (Other
locality—eentra] Vic.)
Gattya trebileocki sp,
FIGS. 48-52.
Tepe Matetlal and Records: Holotype. NMV
(52029, microslide, G2099. preserved mate-
tial, remainder of holotype culony=-R, 10—
33) m, on fragment of Caulerpa brewnil-
paratypes, G2030, G2037, G2032, G2053,
G2034. G2035; SAM H39, microslides—R.
10-33 m, Gn Caulerpa brownit; G2100, pre-
served material, remuinder of paratype colo-
nies,
Description from holorype and paratypes:
Uydrorhiza tubular, of same diam, as stem,
embedded In the stem of the alga, Sreaw to $
mm long, proximal stem intertrode 0.70-0.75
mm long, athecate, perisare thick, with a few
rough annulac constrictions, and 1-2 distal
nematothecue: distal hinge joint V-shaped, Fol-
lowing stem internodes alternately thecale and
athecite, thecale internodes 0.25-0.45 mm
long, athecate internodes 0,10--0.13 mm long.
average diam. of internode 0.10 mm. Hydro-
cladia arising From a short apophysis of the
stem above the bydrotheca on euch cauline
interrode, first pair opposite, following hydro-
cladia altecnate. Hydrocladium with 1-3 hyudra-
thecac, thecate and athecale internodes alter-
nate, crowded. identical to cauline internodes.
Athceate internodes very short. 0.04-0.06 mm
long. 2 between hydrocladial apophysis and
first thecate internode, distal joint oblique, the
second internode with oblique distal node. and
{ median pematotheca; following internodes
alternately alhecate and thecate, athecate inter-
nodes 0.07-0.08 mm long, thecate internodes
21-029 mm long, with oblique proximal and
transverse distal nodes, Ilycrothecue distal on
internode, cup-shaped, U.t4-0.20 mm deep,
perisare delicate. base curved, set well Jown
in hydracladtum; abeauline wall 0.15-0.20 ram
long, convex, thickened to hase of median
nematotheca: adcauline wall shorter, 0.12-0,17
min tong (to end of lobes) not thickened.
almost straight, Afargin sinuated. (20-0.25
mm in diam., with 5 lobes of which 3 wre well
developed, tongue-shaped—1 unterior. peaked.
tising over aperture. 2 posterior, paired, with a
deep sinus between, and 2, paired. in middle of
margin, broad and low in lateral view, hut often
obscure Nemarothecae hithalamic, of 3 types—
1 medion, on each athecate internode (except
HYDROIDS th?
first hydrocladial internode), base slender,
distal cup deep and rather natrow; 4 nemato-
thecae on theeate internodes—! snedian, base
very stout, 0.08-0.10 mm long, distal cup €x-
eavated on adcatiline side, cup 0.04-0,06 mm
im diam,, margin vertical. closely adpressed to
hydrocladium; 2 posterior Jaterals, longer than
medians, 0,08-0.10 mm long, distal cup wide
and shallow, 0,05-0,08 mm in diam., slightly
flattened on adcuuline side, overtopping mar-
ginul lobe of hydrotheca, base slender, on a
pedicel 0,06-0.08 mm long; t small median
suprathecal, similar to nematotheewe on athe-
cale internudes, but smaller, deeply set between
the twin laterals at base of hydrotheca.
Hyedranth with approx. 16 tentacles. Colous—
pale yellaw. Gonotheca absent,
Remarks: G. teebilgocki shows close affinity
with G, aglaopheriaformis Mulder & Trebil-
cock, but may be easily distinguished from, this
species by the shape of the posterior marginal
lohes of the hydrotheca, which in G. trebilcocki
are rounded. The cups of the lateru! nemato-
thecae are also much larger in G. trebilcocki.
Both species occur in the same locality and
over a similar depth range, but G. agiaophe-
niuformisx as presently known is an epizvic spe-
cies, whereas G, trebileock? has been found
only on algal substrate, Both species are Tare.
Pluniularin procumbens Spencer, 1891: £40,
pls. 21-23, figs. 17-25. Bale, 1894: 115,
pl, 5, figs Lt, 125 1914a: 29: 1915: 297,
Briggs, L915: 30S, pl. LO, fig 1,
Records: R, 33 m. epilithic on vertical rock
faces.
Material: One infertile colony 7.5 em high.
Srem thick, fuscicled, growing fram a fibrous
rootstock. Shore hydrocladial internodes with
nematothecue ax described by Bale (19140)
and Briges (1915).
Remarks: Although only 1 specimen was col-
lected, several mature ¢culonies of similar size
and appearance were noted, The colonies arc
amall io comparison with Briggs’ Tasmanian
material and Spencer's material from Port Phil-
lip Bay, Vic. Bale (1914a, 1915) does not give
dimensions of FP. wrocwinbens from the Great
Australian Bight,
Phimularia asymmetrica Bale, 191 4a: 29, pl. 4,
figs. 2, 3; 19152 279,
FIG. 53
Records; Sin. F, 65 m, freegrowing on sandy
bottom
Meresial) (ne infertile colony, 30 ¢m high.
Stem jong, flexuous, branched, strongly {s-
cicled near base. Hydrocladia with 12-15
hydrothecae, hydrocladiai internodes with 5-7
strong septal ridges, Hydretheeae long, adnate,
abcauline wall curving over distally towards
hydrocladium; an indistinct intrathecal fold
sometimes present about halfway aluny thecal
wall. Margin with 2 broad bluntly pointed
jateral Jobes, usually of the same size and
shape, occasionally | lobe much more promi-
nen than the other.
Remarks: The Pearson |. material shows some
variations compared with Bale's microslides uf
-Endeavour’ material from the Great Australian
Bight in the collection of the NMV. The “En-
deavour’ specimens show considerubly more
curvature of the distul hydrothecal abcuuline
wall than the Pearson 1. material, Haye a dis-
lince oblique intrathecal ridge, and a maximum
of 4 septal ridges ia the internode. The mar-
ginal lobes of the ‘Endeavour’ specimens sel-
dom show the pronuunced degree of asym-
meliry inferred from Bale’s figures of P. usyu-
metrica. Furthermore, the margins of the lobes
are reunded, rather than pointed, and each pair
is usually the sante shape; however, as the
hydrathecal margin itself is slightly oblique to
the hydrocladial axis, the lobes appear asym-
metrical when viewed from abeve,
P. asynunetica shows g strong resemblance
to «figures and description of P, Aerrmwipzé
Stechow, 1909 from Japan (Bile I9l4a, p. 31)
and P, #uberert var, elengata Billard, 1913
from the Indo-Pacific revion. The latter is a
small species 2-3 cm high, and the hydracladtal
internodes and hydrothecae appeie to be indis-
inguishable from P. asymmetrica fram Pearson
I, The Jess distinct intrathecal fold and more
symmetrical marginal lobes of (he Pearson I,
specimens tends to bridge the gap between /’.
asymmetrica and Py hertwigi. Possibly all are
geographical variants of the one species.
Although only | colony was collected, the
species was a dominant member of the seafloor
community of the deeper water. Many of the
ulder colonies were almost completely invested
hy wa growth of a pink coloured epizoie zoun-
thid, the weight of which bends the colonies
over to touch the sand,
This is the fourth record of Pi asvanmerrlea;
other records are ulso from the Great Austra,
lian Bight.
Plumularia flexuosa Bale, 1894: 115, pl 5,
fies. 6-10. Mulder & Trehilcock, 1916: 78
{discussiun). Stecbow, 1925: 246. Black-
183
burn, 1938: 315. Shepherd & Watson,
1970: 140.
Pluntalaria pulehella (Bale), ‘Totton, 1930:
221, fig. 58.
Records: R. 27-45 m, on Mvchoden carnosu.
Material: A few infertile stems, Siems to 3 mm
long; internodes Jong, flexuous, nutes tans-
verne, 3 cauline nematothecae on an internode
—?2 uxillar, and 1 proximal, exactly as des-
cribed anu figured by Bale (1894) for P,
(lexuutase,
Reraurks; [ have examined a series of micro-
slides of FP pulchella Bale, 1882, and P,
flexwese in the Bale collection of the NMV.
The stems of P, pulehella are robust, with
several transverse cauline internodal septa, but
have no cuuline nematothecae. Hydrocladia
and hydrothecae are identical in both species.
The yornotheca uf P, putchelle is globular,
with an oblique aperture, and 4 rw of large
internal submarginal weth. In PF. flexuosa, the
gonotheca is clongate, twice as long as wide,
and there are no submarginal teetlr.
Although Bale (1894) clearly distinguished
between the two species, Totton (1930) united
them in P. pufchella on the grounds that “P,
flexaesa . - . appears to fall well withia the
Tange of variation of this species™ «ie. P. pul-
chefla) as the stems of some of his material
were “fine and fcxuous, while others were stout
and straight", His synonymy has since heen
followed by Ralph (1961b) and Millard
(1957).
Stem thickness and the presence or absence
of cavling nematotheeae are frequently un-
reliable specific criteria aniong the Plumu-
Jariinae, but tuking into account the difference
between the gonothecuc (unless sexual dimor-
phism can he demonstrated) it scents best, fol-
fowing Blackhurn (1938), tu regard the two
as distinel species.
Although associvted with a range of algal
substrates in other localities (J.W. unpublished)
P, flexnosa occurred only associated with a
delicate species of the red algal genus My-
chedea (nsually placed under M, carnosa in
herburiu), The growth habit of the trydroid is
unusual and was first poticed by Dr. H. B. S,
Womersley and G, T, Kraft who supplied the
following descciption. “The hydroid infests the
Mychodea frands from «@ very early stage, with
frands less than | cm high showing abundant
hydroid stolohs. The stolons penetrate length-
wise through the outer medulla of the alga.
branching occasionally laterally, and producing
at regular intervals through the cortex the erect,
JEANETTE E. WATSON
polyp-bearing axes. As the Mychodva plant de-
velops.. proliferation of the hydroid stolons in
the lower axis breaks down the algal tissue until
the Mychodey is attached to the substrate only
by a detise weft of hydroid. This may be 2-8
mm thick and a centinetre or more jung, sup-
porting a much branched Mychodea plant over
20 cm long and infested throughout with the
hydroid.”
Plunularis spinuloga Bale, 1882: 42, pl. 15,
fig. 8 I8S84: 139. pl. 12, figs, 11, 12:
1888; 783. Stechow, 1925; 246, Mallard,
1962; 301,
FIGS, 54, 55
Plumidaria spinitese var. spinulosa Ralph,
LY6.: 37, fig, 4. Shepherd & Watson, 1970> 140,
Reeordy: R, 18-30 mm, on Lanrencia elute,
Plecamium angustum, Thyroscyphis mar-
ginatuy and Aglaopitenia plumosu.
Material: Abunctant infertile colonies of 2 few
stems ech. Hydrorhiza wide and flat with
transverse dark markings. Stents to 3 mm long.
intemodes of variable length, 0.17—-0,21 mm,
width at node 0.02-0.04 mm Hydrovladia
arising near middle of short internoues, distally
On long internodes. Hydrothecae 0,16-0.18 mm
deep, abcauline wall strongly convex, Terminal
hydroclaciul spines well developed, varying
from long und sharply pointed to blunt and
barely protruding, past hydrothecal margin,
Netnatathecue identical with typical form, but
pedicels of the median hydrocladial nemate-
thecue show considerable variation in thickness.
Reniarks: The Pearson 1, matertal shows a wide
variation in size of the stem internodes, position
of the hydrocladial apophyses, width of the
hematothecal pedicels and length of the ter-
minal spine. Those hydrotheeae with more pro-
hounced terminal spines arc always larger and
more robust in appearance than those with the
Shorier spines. Because of the variahility of
length of the terminal spine, Millard (1061)
n® longer recognizes the distinction. between the
varicties of P, spintlosa (Le var. pypica
Stechow, 1923 — var. spinulosa Ralph, 1961,
and var. obtusa Stechow, 1923). The present
materi] supports her view.
No correlation was eviden| between stem
type, substrate, or environmental conditions.
Plumularia goldsteini Bale. 14982: AL, pl, 15,
fig. 7: 1884; 157, pl.11, fig, 9,
Records: R. 30 m, on Delisea pitlchra,
Material: A few infertile stems to 3 mm lou
Remiarkiy The specimens conform exsctly to
the description of P. poldstein? by Bale.
HYPROIDS
A new record for $, Ayst. (Other loeality—
Vic.)
Plumularia obliqua (Johnston, 1847), Bale,
I884; 138, pl 12, figs. 1-3, Blackburn,
1942. 108.
ai asad obligna Johnston, 1847; 106, pl. 28,
gL
Records: R, 20 tm, on Melanusiephore fla
bellatn; S, 30 mon Sargassum sp,
Material: A few delicate infertile stems to 4
mm long.
Renwarks; The material conforms to descrip-
lions of PF. oblique hy Bale.
Phonulmia australis Kirchenpaver, 1876. Bale.
I884: 143, pl. 12, figs. 6, 7, pl. 19,, digs.
43, 44.
Plumularia obligue vac. adsirelis Kirchenpauer,
(876: 49, pl. 6, fig, 10,
Reeaus: S$, 14 m, on the seagrass Posidonin
australis.
Material: Luximant infertile colonies, Stems to
4 mm long, srising from a broad fat hydro-
rhiza with transverse dark markings.
Remarks: The Pearson Lo material compares
with Bale’s (1384) description of P. wnsyrelis.
The median nematothecne are, however, not is
deeply exeuvated on the adeauline side nor as
closely adpressed li the bydrocludium as in his
figures. ‘The aaillar monothalamic nemato-
thecat are absent from many stems.
Plumularia epibracteolosa o.sp.
FIGS, 56-60
Type Material and Records: Holotype, NMV
(42046, microslide; G210] preserved mate-
nial, remainder of holotype colony; para-
tvpex, G2047, G2048, G2049, G2050,
G205!; SAM H37, microslides; all material
—R, 50 m, on Sargassum bracteolasye,
Description fren feletype and peralypen:
Aydrorhiza flat, reticular, 0.25 mm wide, with
pegeed borders, radiating from a digitate sto-
lunic plate, Steets monosiphonic, to 2 em long,
perisare thick; proximal internodes toughly un-
dulated, without hydrocladia, nodes indistinct,
following internodes hydrocladiate, 0.36-0.45
min iang, 0.14-1.85 mm in diam., proximal
and distal nodes oblique, V-shaped. Perisarc
smooth externally, internally ridged by 3—+
intemodal sepla—t above, and 1 belaw nore,
1-2 in middle of internode, ridges fewer in
younger parts of stem; either absent or inci-
piently developed in younger stems. Hy«lro-
eludfie alternate, 1 on each slem tnternode, tn
| plane, arising from a shor distal apophysis of
189
the stem, One or two hydrothecae on hydro-
Cladium: hydrocladium beginning with 1, occa-
sionally 2, short proximal athecate internodes
0,09-0,12 mim long, proximal, hode transverse.
distal node slightly oblique; thecale inlernode
0.23-0.27 mm long, socketted into the athecate
internade by a slender joint; hydrocladia below
internode straight, blunt end not projecting be-
yond thecal margin; 4—5 oblique interngdal
sepla dividing internode into segments, 2 below
median nematotheca, sloping opposite ways,
and 2 below the hydrothecu. When 2 hydro-
ibecae are present on hydrecladium, they are
separated by 2 athecale internodes, the first
short, ©.07-0.09 mm long, with transverse
joints, the next 0,12-0.16 mm long, with a
socketted proximal 2nd an oblique distal joint
and Ll median nematotheca; both internodes
without septa. Hydrothecae wide, shallow, cup-
shaped, 0.15-0.17 mm deep, with a Hat base,
set an the 3 strong convexitics of the hydro-
cladium; abcauline wall straight, 0.11-0.13 mm
long, thickened by a continuous flange of peri-
sare extending the entire length of the thecate
imernode; adcauline wall 0.10-0.11 mm tong,
slightly convex, adnate to hydrocladium only
neur base, the remainder joined to the hydro-
cladium by a wedge of perisarc. Margin 0,14—
0.19 mm in diam. (laleral view), sinuated, with
a thickened outwardly rolled rim, the line of
the margin curved down to meet the hydro-
cladium, but the aperture truncated by a deli-
cate transverse shett of pensare extending
across the cup 0.05 mm above adcauline wall.
Neimuothecae all of similar shape and size,
0.06-0.09 mm long, bithalamic, distal cups
shallow, entire, 0.03-0,04 mm in diam.; 2
cauline with slendor bases, 1 in middle of stem
internode, often missing, and 1 axillar; 3 on
thecate hydroclidial internodes—i, median,
hase stout, cup slightly excavated on adcauline
side, 2 laterals below the hydrotheca, bases
stout, cups narrow, standing upright on hydro-
cladium but not reaching top of transverse peri-
sarcal web, One minute mamilliform pore pre-
sent on the shoulder of the hydracladial apo-
physis. Aydranth with approx, 24 tentacles.
Gonothecue large, 1-2 on lower stem on
a short pedicel arising from an old hydroela-
dial apophysis, elongate oval, 138-18 mm
long (including pedicel} maximum diam. 0.72—
1.02 mm (at two-thirds the distance up from
pedicel) perisare delicate, smooth, or slightly
uncdulated, no operculum, top closed hy a thin
convex membrane. Gonophores male, mature.
surrounded by a thin blastestyle Celaur—
stems bright yellow, gonothecac orange.
19%) JEANETTE E. WATSON
Fig, 53.
Figs. 54,55. Plumularia spinulosa Bale. Fig, 54.—Part of a stem with larger hydrothecae and prom-
inent terminal spines. Fig. 55,—Stem wiih smaller hydrothecae and blunt spines.
Figs.. 56-60. Plurwlaria epibracteolosa n.sp, Fig,
ot stem showing internadal septa a
Plumularia asymmeirica Bale. Part of hydrocladium with two hydrothecae,
56.— Whole stem with stolonic plate. Fig. 57.—Part
nd cauline nematothecae. Fig. 58.—Hydrocladium
with two hydrothecae. Fig. 39.—Hydrotheca, dorsal view. (Figs. 46-59 drawn from holo-
type). Fig. 60.—Group of two male gonophores, fram paratype,
Remarks: P. epibracteoloxa is closely allied to
the P. setaceoides group endemic to Australia
and New Zealand, It shows some affinities with
P. excavata Mulder & Trebileock, and with P.
corrugatissma Mulder & Trebilcack, bug is
easily distinguished from both these species by
the structure of the hydrothees and from P.
corrugatissma by its greater overall size.
Stolonic reproduction, common among some
species of the Plumulariinae, has been discussed
by Billard (1904) and Gravier (1971) but has
not previously been reported among the Aus-
tralian members of the subfamily. Many stems
of P. epibracteclosa show various stages of dis-
tal enlongation into a tendril which flattens out
laterally into an embryonic stolonic plate. This
plate adheres to the edge of a nearby algal
frond, sending out hydrorhizal filaments to
form a new colony, the parent stem ‘finally
breaking away. [n one case (holotype imicro-
Slide) # stem has re-attached itself by the dis-
tal end to the same stolonic plate, forming a
closed Inup.
P. epibracteolosa exhibits extreme variation
in development of the cauline internodal septa.
The older stems, distinguished by the thicker
perisarc, are heavily internally ridged, while
the younger stems have either none at all or
show a gradational development between the
two extremes. The presence of internodal septa
has often been accepted as a diagnostic charac-
ter within the Plumulariinae, but the variability
of P. epibracteclasa demonstrates the un-
reliability of this criterion,
HYDROIDS 191
The fronds of the substrate alga Sergusrsune
bracteolosum are seasonal, growing fram Sep-
tember to February (Shepherd & Womersley
1970), P. epibracteolasa must therefore spread
very rapidly in order to form fertile colonies
within a very shurt period. This may account
for the unusual propagation of the colonies by
both normal growth and stolonic reproduction.
the fatter method ensuring spread of the
calonics from one part of an alga to another.
Although the alga, §. bracteoloswn, is also
very common at West I, (Shepherd & Womers-
ley 1970), P. eplbracteolosa Was never recorded
from this locality. At Pearson 1, the alga was
restricted ta a: Jimestoie seafloor in moderate
surge at a depth of 50 m, at a distance of 400
m offshore. The colonies of P. epidracrealesa
accur only on the fronds, whereas Amphixbesia
mlninwe var. pumileides Bale exclusively
epiphytises the harder stems of the alga.
Momolaria meretricit o.sp.
FIGS, 61-64
Type Material and Records: Holotype, NMV
G2n55, microslide; G2102, preserved
material, remainder of holotype colony—
R, 27-30 m, on sponge on vertical walls;
paratvpes, G2054, G2055, G2056, G2057,
G2058, G2059, SAM H38, microstides;
G2103, G2104, preserved material, remain-
der of paratype colonies—S, 18 m, on
sponge on tuck walls,
Description fram jelotvpe and paratypes:
Aydrarhiza tubular. Stems monosiphonic, erect,
straight, to 1S mm long; stem internades 0.42--
0.51 mm long, smooth, the proximal internode
beginning with a transverse joint near hase of
stem, following intemodes with an oblique
proximal joint, often indistinct, and a strong
distal joint, 0.06-0.09 mm in diam. Aydre-
cladie alternate, 1 on each internode, widely
spaced, arising from a distal apophysis 0.05
inm fong. and 0,08 mm in diam, at extremity
of internode, with 1, occasjonally 2 hvdro-
thecve, and rarely, a secondary branch given
Off behind the first hydrotheea, Hydrocladium
with either L long smooth proximal athecate
internode 7.14-0.19 mm long, and 0.07 mm
in diam., of alternatively, 2-3 short athecate
internodes 0,05-0.12 mm long. with internally
Tidged perisarc, These are followed by a long
thecate internode 0.30 mm long, entitely occu-
pied by hydrotheca and an infrathecal cham-
ber 0.10 mm Jong, terminating behind hydro-
thecal margin, Mydrorhecee campanulate, O15
mm deep, at 40" to hydrocladial axis; alcauline
wall rounded in lateral view, almost entirely ad-
nate and immersed in internode, abcauline wall
straight, expanding, contiguous with Iine of
upper wall of hydrocladium, very slightly con-
stricled behind margin. Murgin evérted, 0,18
mm in diam., slightly simuated, curving dows
und back to adcauline wall. Nemutothecae
bithalmic with slender bases and shallow distal
cups partially cut away on the abcauline side,
occasionally 1-2 halfway up stem internode an
opposite side ta hydrocladium, and 1 axillar, 3
present on thecate internode—! median, 0.04-
0,06 mm long. adpressed to the intrathecal
chamber, and 2 laterals below hydrotheca,
0.03-0.04 mm long, distal cup entire. One
very prominent monothalamic mamilliform
pore, with 1, sometimes 2 orifices on short
tubular necks projecting from the top of the
stem apophysis, Gonotleca absent.
Remarks: P. meretricia, like P. epibracteclosa;
shows considerable variability of features
usually regarded as reliable specific criteria
among the Plumulariinae, The younger stems
have a smooth glassy appeurance, with long
stem and hydrocladial internodes, and also have
cauline nemutothecac, The stems seldom retain
their chuline nematothecsac after maturity,
showing only scars where the nematothecae
have dropped off, Since many of the younger
siems show neither scars nor nematothecae,
and in some cases, the axillar nematothecae
have failed to develop as well, the presence or
ubsence of these structures cannot be regarded
as diagnostic of the species, The mamilliform
pore is however, a constant feature of all the
stems, The regenerated athecate hydrocladial
internodes common in some stems, mark the
site of repeated hrenkage and regrowth of the
hydrocladium. In these cases, a short athecate
internode is first added, follawed by an em-
bryenic hydrotheca and infrathecal chamber.
Nematothecue bud off Tater as the hydrotheca
nears maturity. The athecate intemode at this
stage is without internal perisarcal ridges. These
develop ag the lrydrocladium ages.
P. meretricia shows some relationship with
P. flexuosa Bale and P. hyalina Bale, but it ts
much larger and more robust than cither of
these species.
Phimalaria togata o.sp.
FIGS 65-67
Type Material and Records: Holotype, NMV
G2060, microslide—R, 33 m, on Metavonin-
lithen eharoides: paratypes. G2061, G2062,
G2063, G2064, G2065, G2n6, SAM Ha4fi,
192 JEANETTE E. WATSON
WW OZ
wuo-|
0.25mm
Figs. 61-64. Plamularia meretricia n.sp. From holotype. Fig. 61—Whole stem, Fig. 62.—Part of stem
with young hydrocladium and smooth athecate internode, Fig, 63.—Older hydrocladinm
with regenerated athecate internodes. Fig. 64.—Mamilliform axillar namatotheca, en-
larged.
Figs, 65-67, Phumiduria togata n.sp, From holotype. Fig. 65.—Part of stem. Fig. 66—Hydracladium
and hydrotheca,. lateral view. Fig. 67-—Hydrotheca, anterior view, showing aperture.
microslides, C2105 preserved material, re-
mainder of paratype colonies—S, 30 m, on
Meteagonialithan charaides.
Description from holotype and paratypes:
Hydrorhiza broad and flat with transverse dark
markings. Stems short, to 4 mm long, mono-
siphonic, flexuous; 1-3 short proximal inter-
nodes with transverse nodes, following inter-
nodes hydrocladiate, longer, 0,03 mm long,
0.08 mm maximum diam., smooth, proximal
node transverse, distal node V-shaped, a strong
transverse septum above the node. Hydrocladta
short, alternate, 1 on each internode, arising
from a distal apophysis, with 1 very short
proximal athecate internode 0,03-0,04 mm
long, followed by a longer athecate internode
0,19-0,21 mm long, curving very slightly below
base of hydrotheca. Hydrothecae subglobular,
0.20-0.22. mm. high from base to crest (lateral
view). 0,16-0.18 mm wide (front view). ab-
cauline wall rising perpendicular to hydro-
cladial axis, then curving over and hack to
thecal margin; adcauline wall rounded, set well
into hydrocladium, free part rising in a sinuous
curve to the margin. Infrathecal chamber 0,1 1—
0.13 mm long, maximum width 0.10-0.12 mm.
the proximal joimt slenderly pointed and
socketted into the athecate internode. Nernato-
thecae bithalamic, with slender bases, terminal
cups wide, a little cut away on adcauline side:
2 cauline—1 axillar, 0.07 mm long, and 1,
same as axillar, one third distance up internode,
on opposite side to hydracladium; 3 bydro-
cladial nematothecae, 1 median, 0.05 mm tong,
HYDROIDS
adpressed to. the infrathecal chamber. 2, slightly
smaller, standing upright on a projection of the
hydrocladium below hydrotheca, barely reach-
ing thecal mirzin, and separated at the base by
a Tounded prominence. of the hydrocladium.
Gonotheea absent.
Remarks: P. jegata is a very small species
closely allied to P, fivalixe Bale, from which it
may be distinguished by its smaller size, the 2
cauline nematothecac, greater curvature of the
abeauline hydrothecal wall and the distinctively
hooded appearance of the margin.
Plamolaria australiensis o.sp.
FIGS. 6871
Type Materia? and Records: Holotype, NMV
G2067, microslide, G26 preserved mat-
erial, remainder of holotype colony—R. 20-
25 m on sponge: paratypes. G2068, G2N69,
G2070, G2071, SAM H41, microslides—R.
20-25 m, on sponge.
Wescrintian from holotype and paratypes:
Hydrorhiza tubular, embedded in surface of
sponge. Siem monosiphenic, flexuous, to 15
mm long, petisare thick, occasionally heavily
thickened at point of regeneration of a new
atem from the broken butt of an old stem,
Internodes variable in length, G.06—1.5 mm,
nodes transverse, distinct, width at node 0.)4—
0.25 mm. proximal 24 internodes withour
hydrocladia, Internodes with 6-12 cauline
nematothecve scattered in 2 vertical rows in the
same plane as hydrocladias axillar nemuto-
thecac absent, hut 7 nematotheca usually
present on intetnode just above hydrocladial
apophysis; older internodes with fewer nemato-
thecae. Hydraciadiq to 2 mm long, alternate
to subopposite (exceptionally. lower hydro-
cladia may be opposite} directed upwards in 1
plane from a short apophysis of the stem: 1—3
hydrocladia on internode. arising near Lop,
middle. or hase of interaode, but this is
variable; shorier internodes have fewer hydru-
cladia. Hydrocladial internodes alternately
athecate and thecate, the proximal athecate in-
ternode with 1, occasionally 2 nemutothecae:
following athecate internodes 0.27-0,32 mm
long (measured along base of hydrocladium)
with a transverse proximal, and strongly
oblique distul joint, and 2 nemaflothecie.
Thecale internodes 0.18-0.20 mm long. 0.06-
0.08 mm in diam. at transverse (distal) node,
4 maximum of 7 thecate internodes on a hydro-
cladiuim, and frequently, 2 transverse intermodal
septum below pedicel of lateral nematotheca.
Thecate Internude with 4° nematolhecue—!
Tus
median, subhydrothecal, 2 lateral, and | supra-
thecal. Alydrathecae asymmetrical in lateral
view, wider than. deep, scoop-shapéd, set al
abvut 45° to the hydrocladial axis, abcauline
wall straight or very slightly concave and a iitle
thickened, 0,16-0.20 mm long; adcauline wall
convex, 0.13-0.19 mm long, the shallow curve
of the wall contiguous with the base of the
hydrotheca. Margin 0.25-0.3L mm in diam.,
entire, delicate, at an angle of 30° to the hy-
drocladial axis. Nematorhecee bithalamic, all of
similar shape and size. the cauline nemnatothecse
with moderately slender bases, cups shallow,
adcauline wall excavated; 2 median. nemato-
thecae on athecate mlemode, similar to cauline
nemiatothecae, O.07-0.08 mm long, bur with
more robust bases, closely sdpressed to inter-
node, the proximal nemutothecae frequently
somewhat smaller than the distal. Theeate tnter-
node with 1 median subhydrothecal nemato-
theca, cup deeply excavated, pressed close to
base of thecal wall; 2 taterals with shallow open
cups 0.05-0.06 mm in diam... slighthy cnt away
on udcauline side, the edge of cup not reaching
thecal margin, base slender, seated on a pedicel
0.05-0.06 mm long, arising at the junction of
the adcauline wail with internode; 1 small leaf
shaped monothalamic suprathecal nematotheca,
set deep in sinus behind hydrotheca, the aper-
ture facing inwards, Gorothecaa present, male
and female urising beside median subhydre-
thecal nematotheca, usually tn proximal region
of hydrocladium, sexes usually separate, occa-
sionally both sexes present on same hydro-
chidiam. Female gonotheca pear shaped, 0.18-
0.25 mm long {excluding pedicel) 0.42-0,55
mint maximum width, with 1-2 nematothecae
in the basal region similar to the laterals, but
lurger. Operculum a thin flap of same size a5
top of gonotheca. Male gonotheca smaller than
fernale, slipper-shaped, 0,13-0.16 mm wide,
with 1 proximal nematetheca, a little smaller
than those on female ponotheca. No uper-
culum, Pedicel a small round seginent 0.07 mm
in diam, in both sexes,
Remarks, P. catitraliensis is closely related to
P. bedoti Billard from the Indo-pacific and P-.
westnt Jarvis from South and Last Africa, but
is distinguished from both these species hy the
shallow scoop-shaped hydrothecae. It also
shows some affinities with some Indo-Pacific
members of the genus Halepteris, ea. H. buski
(Balc) (a deeper water species common on the
sOuthern Australian constline, alsa found at
Pearson 1.}. and with A. polymorpha (Billard)
in size gnd shape of the gonothetse and nema-
194 JEANEITE
tothecae, general aspect of the colonies, and
the tendency toward opposite branching in the
hasal stem region,
Aglaonhenia plumosa Bale, 1882: 37, pl. 14.
fig. 6; 1884: 153, pl. 14, fig. 5. pl. 07, fig.
12 Blackburn, 1942: 110. Stechaw, 1925:
260. Ralph, 1961b: 65, fiz. 9. Shepherd
& Watson, 1970; 140.
Recards: R, 24-33 m, an ascidians, bryozoa-
and Carpopeltis phyllaphera.
Material: Sparse infertile colonies. Stems to 1
om long.
Remarks: The stems are short, with closely set
hydrocladia and robust hydrothecae, features
characteristic. in this species, of an ocean en-
vironment.
Thecocarpus divaricatus (Busk) var. maccoyi
(Bale. 1884: 162, pl. 15. fig, 7 pl. 17, fig,
7), 1915: 312, pl. 1.
“Avlaophenia maccoyi Bale, 1882: 36, pl. 14,
fig. 2. Blackhiirn, 1942: 110,
Thececarpus divaricates CBusk). Shepherd &
Watson, 1970: 140,
Records: R, 23-45 m; §, 4-12 m, on Meta-
goniolithon charoides, Plocamium cartila-
gine, Acrocarpia paniculata and Zonaria
splralls.
Material; Luxuriant fertile colonies. Srenrs
short, ta 4 cm long, given off in groups from
a winding hydrorhiza, Proximal region of stem
without hydrocladia, lightly fascicled, some af
the supplementary tubes running up the main
stem for u short distance then branching. off.
Aydrocladia 4 mim long. Hydrathecue close-set
at 45° to hydroclacial axis, marginal teeth
deeply cut, the second anterior pair outwardly
bent, the unpaired anterior tooth — well
developed, the hatchet shape becoming mare
pronounced distally ylong the branch, Median
nemidothece vatlable in length, just over-
tapping margin in proximi region af hydro-
eladium, increasing to twice the height of
hydrathecs distally, standing well out fram: the
margin. the terminal aperture af the sume
time broadening out into 2 lobes. Corhulae
immature, with 4-11 pairs of gonohydroctladia;
immature gonophores in corbulae with more
than 10 Ieafiels. Cofour—variable, light to dark
brown.
Thecocarpos divarticatus {Busk) var, brigesi
Bale. 1926: 22. fix. 5.
Aglaophenia divaricatr (Busk).
162, pl. 05, fig. 8, pl, 17, fig. 7,
Balu. 1884:
E. WATSON
Aglaophenic ditaricata vay. acanthocarpa’ Bale,
1915 FIZ.
Records: R, 24-33; 8. 18 m, on fragments
of red alge and Canlerpe simpliciusente:
Stn. F, 65 m, on Syniplectoseyphus subdicho-
TOTTER,
Material> A few infertile stems in each colony.
Srenis to 1.5 em long, unbranched, mono-
siphonic, given olf singly from a winding
hydrorhiza, Hydrocladia flexuous. distant, cach
internode with 2 distinct seplu. Aydrothecar
with 4 pairs of marginal tecth. similar in shape
and size, the median anterior tooth net well
developed. Median nemutoiheca slightly lenger
than hydrotheca, following curve of the ab-
cauline wall, becoming erect just behind mar-
gin. terminal orifice round. in some cases
broadening tnto Iteral lobes; canting nemuato-
thecae larger than laterals, bent around stem,
orifice facing posteriorly, (nfaue—hrown.
Remarks: This is the first record of the vat.
hriggsi from §, Aust, (Other locality. Port
Jackson, N.S.W.).
Thecocarpus divaricatus (Busk} var. cystifera
Bale, 19157 314,
FIG. 72
Records R, 24-33 m, 8, 24 m, epillthle on
vertical rock faces,
Material: Abundant. infertile calonias. Colonies.
of 1-3 stems to 10 cm bigh, growing from a
small Common fibrous rootstock, Stenry thick.
woody. brittle, lightly fascicled, the poly-
siphonic tuhes running up the main stem and
branchng out alternately in one plane, giving
the colony a distinct "front and back” aspect
Proximal region of stem barc, showing sears
where brinches and hydrocladia have dropped
off. Aydrocladia to 13 mm long. Hvdrethecie
set al an angle of 45° to hvclrocladial axis; mar-
final teeth of similar size. evenly spaced. the
sinus between often wide wnd shallow. Median
neinatorheca following curve of the abcaulinc
wall. Lenminating just below margin, terminal
orifice round; cauline nematothccae large, ege-
shaped, Colonr—iight brown,
Remarks: Bale (1915) described; but did not
figure the variety eystifera. distinguishing it
from other varieties of T. divaricatus only on
the presence of the enlarged caine nemato-
thecae,
Remarks an the varieties af T. divaricatus:
Tt is of interest that the 3 varieties of this
species, recognized by Bale, ale recorded for
the first lime fram one Jocality, 7. muceovi
ELYDROIDS.
has previously been reported from various
localities along the Victorian coastline (Bale
1884) and from South Australia (Blackburn
1942. Shepherd & Watson 1970). T. cystifera
has been recorded only from) South Australia
(Bale 1915) and 7. hriggsi only from New
South Wales (Bale 1926). The only informa-
tion hitherto available on the macrostructures
of the hydrocaulus is given by Bale (1884)
who described the typical form as having
“numerous divergent branches and very dark
colour” and the var. meccoyi as a “dwarf
form”. (The larger “typical” form, ic. A,
divaricata Busk, a very common and distinctive
species of the south-eastern Australian coasi-
line, was not found al Pearson [., despite
careful search). The distinction between the
varieties has therefore Jargely rested on micro-
structures alone.
195
Although some intergradation in structure
does exist between the varieties, the material
from Pearson I. now enables a clear distinction
to be made in both micro- and maero-siruc-
lures, as Well as environmental preferences. T-
maccoyi and T, brigyysi, because of their simi-
larity in size and overlap ot substrate prefer-
ences, are difficult to dlistinguish in the field, but
they are easily separated on micro-structures:
T. cystifera although unmistakable im size and
growth habit, has hydrothecae almost identical
with those of I. briggst. T. maccoyi was the
only Variety fertile at the time of collection.
The gradation in micro- and macro-structures,
habit, and apparent difference in fertile season
of the varieties, suggests incipient speciation
within the 2. divaricamus group, Distinguishing
features between the varieties from Pearson I,
aré tabulated below.
T. maceeyi
medium, 4<¢m
lightly fascieled,
branched
twice length of hydra-
theca, orifice lobed
Stem length
Colony
Mesial nematotheca
Cauline nematotheca normal size, shape
Marginal teeth sharp, deep
Habit epiphytic
T. cystifera
large, 10cm
T. briggst
small, 1.5 cm
unfascicled, unbranched
to hydrothecal margin,
orifice round to lobed
Fascicled. braiched tn }
plane
to hydrothecal margin,
orifice round
normal size, shape, large, ovale
facing posteriorly
sharp, deep wide, shallow
epiphytic-epizoic epilithic
Lytnearpus mulderi (Bartlett, 1907),
FIG, 73
Aplaophenia muldert Bartlett, 1907; 66, Mulder
& Trebilcock, 1916: 73, pl, 10, fiz. 3-
Records: Among algae; no other data re-
corded.
Material: A fragment | em long, the distal end
of a fertile stem. The specimen conforms. to
vescriplions of Bartlett und Mulder & Trebil-
cock. Gonosome comprising 2 gonophores—t
male and 1 female, in an open corbula arising
from a primary hydrocladium. Primary hydro-
ciasdium with thecate proximal internode, fol-
towed by a swollen internode hearing 3. nema-
tocladia and gonophores. Nematocladia 0.75—
(.84 mm long (but may be broken) each bear-
ing a single row of nematothecae. Ganothecae
round, laterally compressed; female, 135 mm
in diam., slightly larger than male, packed with
mature ova, blaslostyle almost filling ganothecal
cavity: male gonophore surrounded by a blasta-
styit o£ the samé shape, but of smaller size
than the female.
Remarks; This is the first record of a species
referahle 10 the genus Lytecarpuy from
southern Australian waters. As earlier descrip-
tions wete derived from fragmentary infertile
material, it was assumed, in the absence of the
gonosome, to belong to the closely related
genus Apglaophenia, common in southern Aus-
tralia, This is the vhird record of this rare but
distinctive species, and the first record for S,
Aust, (Other locality—Bream Creek, Vic.).
Halicornopsis elegans (Lamarck, 1816). Bale,
tda: 56; 1915: 303. Briges, 1914:
309, Blackburn, 1942; 107. Shepherd &
Watson, 1970; 140,
Plamiularia elegans Lamarck, 1816; 129,
Halicornopsis aviculuris Bale, 1882: 26, pl. 13,
fig. 3; 1X84: 185, pl. 10, figs. 1. 2. pl 19, fig.
32.
Records: R, 33 m, epilithie, and on bryozoa
and red algae.
Material: One small infertile colony. Stems Lo
3.cm long. branched,
Remarks: Vhe colonics were comparatively
small and the individuyl stems short for the
species,
JEANETTE E. WATSON
196
O.5mm
wus 'o
wusz:-O
wuUgrd
wu Q-2
HYDROIDS (97
Halicornaria longirostr® (Kirchenpauer, 1872).
Bale, 1884: [8]. pl. 13, fig. 7, pl. 16, fig.
3, pl. 19, fig. 30. Shepherd & Watson,
1970; 140.
Apgloophenia flongirastris Kirchenpauer,
28, pl. 1, fig. 19, pl. 5, Ng. 20,
Records: KR, 18-33 m, cpilithic on rock walls,
oo Herainunia memus, red algae, and bryo-
zoa,
Material: Abundant infertile colonies, Stems to
7 om long. with 1-2 proximal branches. Colour
—pale straw colour
Remarks! The colonies fall into 2 groups—
those with long stems, and those with short
stems, The larger colonies, comprising clusters
mt longer stems (up to 7 cm) were either epi-
lithic or epizoic, whereas the shorter stems Cup
tu 3 cm) growmg singly, were epiphytic on
algac. There is no difference in micro-structures
between these two ecologically distinc, stem
types.
Halicornaria prolifera Bale, 1882: 34, pl. 14,
fiz. 5; 1884: 183, pl). 14, fig. 1, pl. 16, fig,
10, Ritchie, 1911; 858, pl, 85. figs, 2, 3.
Records: R, 30 m, epilithic.
Marerials One infertile unbranched colony.
Stem & cm high, Aydrocledia 0.7 mm long.
given off at un acute angle to the stem. Anterior
and posterior cauline nemutothecae with 3,
somerimes 4 orifices, median hydrocladial
nematotheca extending just below thecal mar-
gin. Marginal teeth shallowly scalloped, the
middle pair slightly everted.
Remarks: The specimen agrees in most respects
with Bale’s description of H. prelifera, except
that the median nematothecue ae a little
shorter than those described by Bale, and all
the cauline nematothecae have 3 orifices. The
hydrothecal margin is circular in anterior view,
similar to Ritchic’s (1911) specimens. This is
the first record ol H. prolifera from S. Aust,
(Other localities—N-S,W. and Vic.)
Halicornaria auren 1sp-
1&72:
material, remainder of holotype colony—R,
33 m, epilithic on rock walls; paratypes,
Gi089, microslide, G2108, preserved
material, remainder of colony—R, 33 m,
epilithic; G2090, microslide, G2109, pre-
served material, remainder of colony—R,
27-30 m, epilithic on rock walls; SAM
microslide.
Dexeription from holotype: Colony 6 em high,
growing from a smail fibrous rootstock, Sent
monosiphonic, lower stem 1 min in diam,, athe-
cute. divided inte internodes, nodes transverse,
proximal intemodes wath circular pits where
cauline nemuiethecue huve dropped off. First
branch 2 cm abuve base, all branching there-
after dichotomous, at an angle of about 40°,
the branches becoming somewhat convergent
distally, then rebranching, Branching repeated
6-7 Times, always in the une plane. Branch
internodes short, 0.60-0.69 mm, divided hy
indistinct transverse nodes, diam, at node,
0.66-0.84 mm. AHydraciadi2 to 5 mm long,
alternate, 2 on an internode, given olf after
first branching of main stem, standing out
stiffly at an acute angle from the branch, giving
the colony a decidedly “front and back” aspect.
Hydrocladial internudes 0.28-0.31 mm long,
nodes almost perpendicular to the axis, indis-
tinct, no internodal septa, Hydrothecae squat,
set. at an angle of 50° to hydrocladium, 0.23—
0.26 mm deep, filling internode; adcauline wall
straight, fixed part 0.12-0.14 mm long, free
part 0.03-0.05 mm long; abcauline wall 0,t7—
0.19 mm long, divided in the middle by a long
intrathecal ridge projecting slightly forward
more than half way across thecal cavity; base
of bydrotheca flat, with a small knot of den-
ticles on the adcauline side marking the hydro-
pore, Margin (.17-),29 mm in diarm., with 8
teeth—3. pairs of prominent blunily pointed
teeth, the middle pair the longest, outwardly
hent, the anterior pair erect. the posterior pair
slightly everted: 1 low postenor tooth, often
chsolete. and 1 small anterior tooth below the
median nematotheca. Median nematothecue
almost twice the height of the hydrotheca,
0.28-0.40 mm jong, 0.05-0.07 mm wide at
FIGS. 74-76
Type Moterial and Records; Holotype, NMV
G2088, microslide; G2107. preserved
Figs,
#8-71. Plumularia australiensis usp. Fig. 6%.—Part of stem. Fig, 69.—Part of hydrocladium, en-
larged. Fig. 70.—Male gonotbeca, (Figs. 68-70 from holotype), Fig, 71—Female eonp-
theca, from paratype.
Fig. 72
to shaw cauline nematothecae.
Fig 73.
Figs. 74-76
Thecoéarpus divaricatus var, cystifere. Bale. Part. of branch with hydrocladia removed
Lytocarpus mulderi (Bartlett), Open corbula with male and female gonophores.
. Halicorseria aurea op. From holetype, Fig, 74.—Part of stem wilh hydrocladia on one
cide removed ta show cauline nematothecae, Fle, 75. —Hydrothecae. anterior view, Fig,
76,.—Hydrothecsa¢, Jatersl view-
198 JEANETTE EB, WATSON
hase, tapering distally and inclined forward,
lerminal apeyture small, circular, lateral aper-
ture distinct, Lateral neiatothecae small, 0,1 1—
O13 mm long, saccate, not reaching thecal
margin, | small terminal aperture on aw short
outwardly turned neck, and 1 lateral aperture
facing inward towards the hydrotheca. Gono-
theca absent. Colaur—amber,
Remarks: The matginal thecal teeth exhibit the
variations in Jength and shape characteristic of
Halicorrnaria. The teeth are normally long, the
middle tooth being the longest of the 3 on each
side. The median nematothecue are all of
newly equal size, and show little tendency
towards increase in length in the distal region
of the hydrocladium.
A. aurea resembles 2 other southern Austra-
lian species of Hallcornaria—H. superba Bale,
and #f.. baileyi Bale. Tt differs from the former
in minor micro-structures, the marginal thecal
teeth of H. superba beng sharper and narrower
than those of I, aurea, the median nematu-
theca of H. superba is larger, and the lateral
nematothecae have 1 Jateral and 2 terminal
uperiures. In A. aurea the Ioterals have only
2 apertures, i facing inward and the other out-
ward; as they are very small they are sometimes
difficult to distinguish, In mucroe-structures,
however, H, aurea is easily distinguished from
Hi. superba, whose stems are long, gracefully
pluiyosc, and yellow-green in colour. In size,
growth hubit, and colour. the colonies af H.
area are indistinguishable from AH. baileyi,
‘they are however, quite different in micro.
structures,
ff, aurea is an abundant species on tock
faces exposed to surge.
Acknowledgments
Jam grateful to the Royal Society of South
Australia and the Department of Fisheries und
Fiuuna Conservation of South Australia for the
Opportunity to participate in the Pearson Island
Expeuition; to the Director and Dr. B. J. Smith
of the National Museum of Victoria, Mel-
boume. Jor use of facilities and equipment; to
Dr. H. B.S. Womersley, Botany Department.
University of Adelaide, for identificaGon and
information on algne, and helpful.advice; to the
Australian Museum for the loan of material,
and to Mr. S.A. Shepherd for criticisin of the
manuseript. Thanks are due to Normalair Gar-
fet, Melbourne, for the loan of a compressor.
L especially acknowledge the unfailing help and
enthusiasm of my diving companions, Scoresby
A, Shepherd, and John O, Ottoway. Expenses
of this study were partly met by a gruna fram
the Science & Industry Endowment Fund,
CSIRO,
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Aust. 97(3), 201-205, 31 August, 1973.
Three species of opisthobranch molluscs from Pearson I., Notarchus indicus (Schweigger),
Aporodoris merria Burn n.sp. and Goniodoris meracula Burn, and one species from nearby
Flinders I., Sagaminopteron ornatum Tokioka & Baba, are new records for South Australia.
Additional distributional data are given for G. meracula and S. ornatum.
PEARSON ISLAND EXPEDITION 1969+—10. OPISTHOBRANCHS
by Rogert Burn*
Summary
Burn, R. (1973).—Pearson Island Expedition 1969.—10. Opisthobranchs. Trans. R- Soc. 8.
Aust. 97(3), 201-205, 31 August, 1973.
Three species of opisthobranch molluscs from Pearson I., Notarchus indicus (Schweigger),
Aperodoris merria Burn usp, and Goniodoris meracula Burn, and one species from nearby
Flinders I, Sagaminopteroa ornatum Tokioka & Baba, are new records for South Aus-
tralia. Additional distributional data are given for G. meracula and S. ornatrm.
Introduction
The opisthobranch molluscan fauna of South
Australia is relatively poorly known, especially
with regard to the naked or “nudibranch”
species Of the western coastline of the State..
Thus it is not unexpected that any collection
from this atea should contain new species and
new records,
Three species described in this paper were
collected during the 1969 Pearson Island Ex-
pedition, organized by the Royal Society of
South Australia and the South Australian De-
partment of Fisheries and Fauna Conservation.
The Pearson Islands are the small southern
part of the Investigator Group at the eastern
end of the Great Australian Bight. The largest
island, Pearson I, is about 162 hectares in
extent, but the others are much smaller; they
lic 64 km southwest of Elliston.
The fourth species im this paper was
collected during a stop-over by the Expedition
at Flinders T., a large island that forms the
northern part of the Investigator Group.
The specimens have been deposited in the
National Museum of Victoria, Melbourne
(NMV).
Order CEPHALASPIDEA
Superfamily PHILINACEA
Family GASTROPTERIDAE
Sagaminopteron ornatum Tokoika & Baba,
1964: 218. Bennett, 1966: pls. 1, 120b.
Baba, 1970: 47.
FIGS. 1-2
Material: Flinders I, Jan, 1969; 2 spect-
mens from 10 m on C'ystophora on vertical
rock face in fair surge (NMV. F27399).
The living animals were reported as “hright
blue under water, parapodia edged with bright
yellow”. They are now colourless and 12 and
9 mm long, 6 and 5 mm broad. When alive,
the larger animal was almost 25 mm long.
The one major difference between the
Japanese type specimens and Australian
material (Fig. 1) is the presence of a relatively
large strong shell in the mantle cavity beneath
the posterior caudal lobe of the body. The
shell of the larger Flinders J, specimen (Fig. 2)
is 2 mm Jong by 1.4 mm wide, and is white
with a yellowish transparent protoconch, the
interior of which is open. The shell of a Lord
Howe I, specimen is 2.4 x 1.7 mm, one from
Cockburn Sound, Western Australia, is 2.1 x
1.6 mm, and another from Heron I.. Queens-
Jand, is almost 4 mm long,
Tt must be assumed that the shells of the
Japanese types dissolved during fixation and
preservation.
Though hitherto unrecorded, Sagaminap-
teron ornatum 3s ia fairly common species in
eastern and southern Australia. Specimens
examined by the writer are:
1. Lord Howe 1L—Middle Beach, Jan. 1959,
R. R. Rlackwood, 1 specimen (NMY,
F20717).
2. Queensland.—Heron 1., Capricorn Group,
Aug. 1965, I. Bennett, | specimen. Humpty
* Honorary Associate, National Museum of Victoria, Melbourne, Vic. 3006.
+ Other accounts of the geomorphology and biology of the Pearson Islands are given in
Volume 95, Part 3 (1971) of the Transactions, as wel] as in the present Part,
22 R. BURN
L, Keppel Group, Supt, 1969, N, Coleman, These specimens were collected from be-
1 specimen. tween the intertidal zone and 20 m depth,
3. New South Wales —Angourie, Aug. 1966, from heneath boulders, from brown algae,
A, A. Cameron, 1 specimen. Minnie or from grey branching sponges. They have
Waters, Jan, 1963, G. Biddle, 1 specimen. often been observed, in rock pools and deeper
Bawley Point, near Ulladulla, Dec. 1963, water, to swim with a rapid synchronous un-
f. Bennett, 3 specimens. dulution of the parapodia.
4, Victoria—Bear Gully, Waratah Bay, April
1970, f. Marrow, 6 specimens. Order ANASPIDEA
5, South Australia—Port Noarlunga, Feb, Family APLYSHDAK
1966, R, Balfour, 1 specimen (SAM, Notarchus indicus Schweigger, 1820. Bergh,
14888). Anxious Bay, Dec. 1968, T- 1902: 349. Engel, 1936; 113. Eales,
Castle, 1 specimen. 1944: 12.
6, Western Australia—West of Carnac |,, FIGS. 3-6
Cockburn Sound, Feb. 1971, B. R. Wilsan Material; Pearson [., Jan. 1969, 1 specimen
& N. Coleman, 4 specimens (WAM, 474- from algae on horizontal faces at 26 m
71). (NMV, F27401).
Figs, 1-2. Sagaminopteron ornatum. Fig. 1— Dorsal view of a 20 mm long specimen from Port
Noarlunga, S. Aust., from a sketch by Dr Helene M, Laws, Curator of Marine Inverte-
brates, South Australian Museum. Fig, 2—Shell from larger Flinders I. specimen, dimen-
sions 2 x 1.4 mm.
Figs. 3-6. Notarchus indicus. Fig. 3.—Diagram of mantle cayity and aperture (a—anus, b—dgill,
c—pigment gland, d—mantle apertute, e—opaline gland), Fig. 4—Radular teeth, Fig. 5,—
Jaw element. Fig. 6—Male copulatory organ (p—penis, r—retractar muscles, s—sheath,
t—seminal groove).
OPISTHOBRANCHS
The colourless preserved specimen is 5 mm
jong, 3,5 mm wide and 3 mm high, Fhe head
and neck are invaginated into the smooth vis-
ceral hump. The 1 mm long mantle aperture
(Fig, 30), of che usual external shape, is placed
well forward, The large mantle cavity, not yet
pervious with the dorsal mantle aperture, con-
tains a small lobulated gill (Fig. 3b), a Jong
loop of the intestine terminating well behind
the gill stem at the anus (Fig. 3a), a long
curved granular pigment gland (Fig. 3c}, and
a small white opaline gland (Fig. 3e). No
genital groove is apparent, Short stumpy
thinophores with deep lateral grooves project
fram the head, and a small rounded knob-like
oral tentacle occurs ether side of the mouth.
The jaw elements (Fig. 3) are narrowest
just below the serrulate distal end. The hyaline
radula (Fig. 4) is almost 2 mm long with 21
rows of 20.1.20 teeth. The rhachidian tooth is
slender with sharply pointed cusp and a short
denticle each side. The lobulate first lateral
tooth has onc inner and two outer denticles,
and the next six teeth have op to lwo inner
and four outer denticles, The succeeding teeth
have longer cusps with up to five inner and
nine outer denticles at the third or fourth tooth
from the margin, The murginal tooth has one
innee denticle well back from the tip and
about four incipient outer denlicles,
The minitile male copulatory organ (Fig.
6) agrees exactly with the figure for Mauritius
specimens given by Engel (1936, p. [16, fig.
4) with the exception that the grooved smooth
penis hys not as many spiral turns.
Discusion: The descriptions by Bergh (1902)
and Engel (1936) of specimens From Mauri-
tius, plus that of a specimen from Zanzibar
(Eales 1944), satisfactorily diagnose Nofar-
chax indicus, The spiral unarmed penis justi-
fies the identification of thts very small Pear-
son T. specimen. The smal) number of lateral
teeth ¢20), the marginals of which are not
slender and smooth, does not agree with the
above three descriptions where the lateral teeth
number 43-45, 33 and 32 respectively and the
marginals are long: slender and smooth. How-
ever, the Pearson I. specimen is probably a
juvenile in which the radula has not attained
the full complement of teeth nor the penis the
full number of turns,
Notarchus indicus has a wide Indo-west
Pacific tropical and subtropical distribution,
and hag been recorded from Sydney Harbour,
New South Wales (Engel 1936, p. 119}. It as
w new record for South Australia,
205
Order DORIDACEA
Tribe CRYPTOBRANCHIA
Family DORIDIDAE
Aporodorls merria n.sp.
FIGS. 7-11
Material: Pearson J., Jan. 1969, | specimen
(holotype) from red algae at 52 m (NMV,
F27402).
The alcohol preserved specimen (Fig. 7) is
dull orange-fawn in colour. It measures & mm
long, 5 mm broad and 3.5 mm high. The
notum is covered with various sized tubercles
(Fig, 86), the largest. of Which are somewhiut
flay clavate and up te 0.6 mm in diam, All
tubercles have projecting angles both laterally
and dorsally. Bundles of spicules strengthen
each angle. The rhinophorsl cavities are pro-
tected by four tubercles; a large one ot each
side, a sniall one in front and a small one be-
hind each cavity (Fig. 8a). The branchial
cavity has nine or ten lappet-like tubercles of
various sizes along the margin (Fig, 8¢); these
tubercles are Up Lo 1 mm tong and 0.8 mm
wide and have small projecting angles or
Points on the onter or dorsal face only.
The thick fleshy hyponotum is narrower than
the foot (Pig. 9), from which it is separated
laterally by little more than the foot margin.
The genital aperture opens in the middle width
of the hyponotum. The lamellate Thinophores
are completely withdrawn. There are five mul-
tipinnate gills. The head (Fig, 10) lies within
a deep concavity of the anterior hyponoeuim,
with a grooved ridge-like oral tentacle at each
side of the mouth. The broad font is anteriorly
truncate with the upper lamina leading into
the head cavity where it. is notched, The tail is
thin and broadly rounded.
The thick labial cuticle ts smooth. The
radula is 1.7 mm long and 1.4 mm wide. It
has 40 rows of 41 teeth per half row, All teeth
are hook-shaped und bear a single outer
denticle beside the cusp, except for the two
marginals which have If or more comb-like
denticles (Pig. 11).
The brittle genital organs could not be
examined satisfactorily, The whitish sompulla
is long and winding. The yellow vas deferens
is short and twisting; and terminates in an un-
armed penial sheath without penial papilla.
Discussion: The angular, flattened, clavate
tubercles of the notam and the single promin-
ent outer denticle of the lateral radular teeth
separate A. nrerria from other species of
Aporodoriy Ihering, (886, The unarmed penis
of the new species is similar to that of THor-
204
disa Bergh. 1877, but in that genus the Jateral
teeth are smooth. The radula is also similar to
that of Taringa Marcus, 1955, but the un-
armed penis of A, #erria contrasts with the
cuticularized penial papilla and spines of the
former genus.
The concavity of the head and the large
lappet-like tubercles of the branchial margin
ure further distinguishing characters, The rela-
tively flat underside of the specimen, with the
head parts recessed, suggests that A. mierria
has unusual feeding preferences.
The specific name is derived from “merri”,
an Australian Aboriginal word meaning stones,
in allusion to the notal tubercles.
Figs.
R. BURN
Tribe PHANEROBRANCHIA
Superfamily SUCTORIA
Family GONIODORIDIDAE
Goniodoris meracula Burn, 1958: 27; 1966:
227,
FIGS, 12-13
Material: Off Dorothee, Jan. 1969, 1 speci-
men [rom algae at 65 m (NMV, F27400).
The colourless preserved specimen measures
7.5 mam long and 3,5 mm broad. Living speci-
mens (Fig. 12) are usually yellowish with
darker brown mottling. Important characters
for the identification of this species are the
smooth body, the high notal flange open be-
ra
lateral =
7-11. Aporodoris merria. Fig, 7—Dorsal view of preserved holotype. Fig. §—Notal tubercles
from rhinophoral cavity (a), middle of the notum (b), and branchial margin (c). Fig.
9.—Anferior hyponotum. Fig, 10.—Detail of head with anterior foot folded down, Fig,
11.—Radular teeth,
Figs. 12-13. Gonivdoriy meracuta. Fig. 12.—Dorsal view of an $ inm long specimen from Point Dan-
ger, Torquay, Vic. Fig. 13—Half row of radular teeth from Sydney Harbour specimen,
OPISTHOBRANCHS
hind the gills, the short caudal crest, and the
seven gills.
The species has been very rarely collected.
The holotype was found eating into a yellow-
ish compound ascidian beneath a. stone at Point
Danger, Torquay, Victoria (Burn 1958), and a
second specimen was recorded from Portsea
Pier, Port Phillip Bay (Burn 1966). A third
specimen was taken by the writer at Point
Danger, Torquay, Dec, 1963, where it was
crawling on brown algae.
Three specimens (Australian Museum,
C312), dredged in Sydney Harbour, New
South Wales, on |! June 1892, are a new
record for that State and the only other speci-
mens known to date. Each measures 10.5 mm
205
long by 5.5 mm wide. The radula (Fig. 13)
of one specimen has the formula 26 x
1,1,0.1.1; the lateral tooth is strongly hooked
with smooth cusp, while the marginal tooth
has much the same shape and is about half.
the size of the lateral tooth.
Goniodoris meracula is a new record for
South Australia.
Acknowledgements
The writer is indebted to Mrs Jeanette E.
Watson, Honorary Associate, National
Museum of Victoria, Melbourne, for the col-
lection of, and the ficld notes on, the Pearson
I. opisthobranchs. This research has been aided
by a grant from the Science and Industry En-
dowment Fund, C.S.I.R.O., Canberra.
References
Baba, K. (1970) —List of the Gastropteridae and
Runcinidae of Japan. Collecting and Breed-
ing, 32(2), 46-48 (in Japanese).
Bennetr, EL (1966).—"“The Fringe of the Sea.”
(Rigby: Adelaide, )
BercH, R. (1902).—Malacologische Unter-
suchungen. Jn C. Semper (Ed.), “Reisen im
Archipel der Philippinen", Vol. 7, pp. 313-
382, plates 25-29.
Burn, R. (1958).—Forther Victorian Opistho-
branchia. J. Malac, Soc, Aust. 1(2), 20-36.
Burn, R. (1966).—Port Phillip Survey 1957-
1963: Ophbisthobranchia. Mem. natn. Mus.
Vict. 27, 265-288.
Eaues, N. B. (1944).—Aplysiids from the Indian
Ocean, etc. Proc, Malac. Soc. Lond, 26, 1=
ENGEL, H. (1936).—Some additions to our know-
ledge of the genus Notarchus, Proc. Malac.
Sac. Lond, 22, 113-119.
Toxioxa, T., & Basa, K. (1964).—Four new
species and a new genus of the family Gas-
tropteridae from Japan. Publs Sete Mar,
Biol. Lab. 12(3), 201-229, pls. 10-13.
PEARSON ISLAND EXPEDITION 19697-11. CRUSTACEA: ISOPODA
BY W. F. SEED*
Summary
SEED, W. F., (1973) .-Pearson Island Expedition 1969.-11. Crustacea: Isopoda. Trans. R.
Soc. S. Aust. 97(3), 207-212, 31 August, 1973.
Eleven species of isopods, all sphaeromatids, are represented in a small collection from Pearson I.
Of these species, two are too immature for specific identification, but belong to the genera
Exosphaeroma and Cymodopsis. One new species, Cilicaeopsis floccosa, is described and figured.
The previously-known species are: Cymodoce gaimardii, C. pubescens, C. unguiculata, Cilicaea
latreillei, Cerceis acuticaudata, Haswellia anomala and H. cilicioides.
PEARSON ISLAND EXPEDITION 1969;—11. CRUSTACEA; ISOPODA
by W. F, SEED*
Summary
Seep, W. F., (1973}.—Pearson Island Expedition 1969.—11. Crustacea: Isopoda. Trans. R.
Soe. §. Aust, 97(3), 207-212, 31 Angust, 1973.
Eleven species of isopods, all sphaeromatids, are represented in a small calection from
Pearson I. Of these species, two are too immature for specific identification, but helong to the
genera Exosphacroma and Cymeodopsis. One new species, Cilicaecpsis floccosa, is described
and figured. The previously-known species are: Cymodace gaimardii, C. pubescens, C. ungui-
culata, Cilicaea latretilei, Cerceix aculicaudata, Huswellia anomala and H, cilicioides.
Introduction
This paper discusses the isopod crustaceans
collected during the Pearson I. expedition of
6-13 January, 1969, sponsored jointly by the
Royal Society of South Australia and the De-
partment of Fisherics and Fauna Conservation
of Soutb Australia. For discussion of collecting
siles see Shepherd & Womersley (1971) and
Watson (1973); R and § indicate rough-water
and sheltered localities.
All specimens Were recovered during the
sorting of algal collections made by divers
(8. A. Shepherd, J. EK. Watson and J. Otta-
way), This may account for the immaturity of
much of the material, smce larger, and pre-
sumably more vigorous, animals are often ob-
served to escape the net during collection
(S. A, Shepherd, pers, comm.). Nine species
(including one new species) are identified, and
two Species are diagnosed to genus only,
The keys of Hale (1929), Hansen (1905)
and Hurley (1961) were used to identify the
genera; most of the speciés were determined
fram Hale’s (1929) keys and species descrip-
tions. Syhonymies ute not necessarily com-
plete. | have followed Menzies (1962; see also
Menzies & Frankenberg 1966) in giving only a
bricf diagnosis of the new species, supported
by accurately drawn figures.
Use of the name Sphaeromatidae (rather
than the more commonly-used Sphueromidae)
follows Schultz (1969) and Naylor (1972),
and anticipates a forthcoming paper by Hurley
& Jansen (pers. comm.) in which the usage ts
discussed,
Specimens are deposited in the isopod cal-
lection of the National Museum of Victoria,
Melbourne (NMV), but only the new species
has been registered.
Tribe FLABELLIFERA
Family SPHAEROMATIDAE
Group HEMIBRANCHIATAL
Genus EXOSPHAEROMA Stebbing, 1900
Exosphaeroma sp-
Locality: Pearson I. (Station F at 65 m),
Material: One immature male, damaged
(about 5 mm long). Penes are developing
but appendix masculina is not yet distinct.
Remarks: More than thirty species have been
placed in this genus, and the specimen differs
from all of them in at least one point: the
uropodal exopod is W-shaped at the distal end,
Description of this species must await more
suitable material.
Genus CYMODOCE Leach, 1813-14
Cymodoce gaimardii (Milne Edwards). Han-
sen, 1905: 121, Baker, 1926: 256, pl,
42, fig. 2. Hale, 1929: 286, fig, 284.
Nierstrasz, 1931: 200. Naylor, 1966: 186.
fig. 2.
Sphaeroma gaimardii
1840; 209,
Milne Edwards,
“Department of Applied Biology. Royal Melbourne Institute of Technology, 124 fa Trobe
St.. Melbourne, Vic. 3000.
* Other accounts of the geomorphology ard biology of the Pearson Islands are given in
Volume 95, Part 3 (1971) of the Transactions, as well as lhe present Part.
208
FIGS, 1-4
Localities: Australia (Milne Edwards 1840);
Vic.: Port Phillip Bay (Baker 1926, Nay-
lor 1966); Tas,: (Baker 1926); S. Aust:
Encounter Bay, Gulf St. Vincent (Baker
1926), New record: Peurson |. (Station F at
65 m).
Material; One female (11 mm long), im-
mature, With no oostegites or eggs, and the
mouthparts unmodified.
Remarks: The specimen agrees with the des-
criptions and figures of Baker (1926) and
Naylor (1964), allowing for is being a
juvenile. Comparison with these figures and
with specimens collected from Western Port
Bay reveals variation within the species in the
sharpness of truncation of the uropodal endo-
pods, in the shape of the joint between telsonic
and pleonic tagmata, and in the shape and
degree of exposure of the anterior suture on
the pleon (Figs. 1-4).
Cymoaduce pubescens (Milne Edwards). Han-
sen, 1905; 122. Stebbing, 1910: 104,
Nierstrasz, 1931: 198. Naylor, 1966:
188, fig. 3.
Sphaeroma pubescens Milne Edwards.
1840; 209,
Paracilicuew (2) pibescens (Milne Ed-
wards), Baker, 1926: 262, pl. 43, figs.
8-11; pl. 48, fig. 1. Hale, 1929: 290_
Locatlties: Austrilia (Milne Edwards 1840);
WN,S.W.: Port Jackson, Port Stephens (Has-
well 1882); Vie.; Port Phillip Bay (Nuylor
1966), Zanzibar: Wasin (Stebbing 1910),
Indonesia: Sailus Besar, Paternoster 1.
(Nierstrasz 1931), New record: Pearson 1,
(Stution F at 65 m),
Material; One mule (8 mm jong) and one
female 7.5 mim long), both immature: the
W. F. SEED
appendix musculina of the male is not lree,
the female has no oostegites Or eggs, and its
mouthparts are unmodified.
Remarks; Specimens agree with published des-
criplions of this species. It should be noted
that, although Hale (1929) has followed Baker
in referring this species to Purarilicaea im the
text, his key agrecs with those of Hansen
(1905) and Hurley (1961), the species key-
ing out to Cymedece in all three.
Cymodoce unguiculata Barnard, 1914: 394, pl.
34B. Baker, 1926; 259. Hale, 1929: 285.
Localities: South Africa (Barnard 1914),
S, Aust.; Beachport (Baker 1926), New
record: Pearson I, (Station F at 65 m).
Material; Two feniles (6 and 9 mm long),
both immature, without eggs or modification
of the mouthparts, but with four pairs of
oostegite buds in the larger specimen,
Remarks: The specimens Jack the marginal
fringe of setae referred to by both Hale and
Barnard; the bosses on the telson are much
less prominent than Barhard's figures indicute
(by inference from his description, and
directly from Hale's. they are the same size in
adults of both sexes); and the hooked uro-
podal exopad differs slightly in both specimens
from Barnard’s figures, The uropodal endo-
pod, however, is. slightly excavate distally, as
shown in Barnard’s figure of the female, and
in other Tespects the specimens agrée well with
desenptions of C. unguiculaia.
Genus CILICAEA Leach, 1818
Cilicaca curtispina Haswell, 1881b: 185, pl. 3,
fiz. 4. Stebbing, 1905: 36, Baker, 1908;
142, pl. 4, figs. 12-17, pl. 5, figs. 1-%;
1938: pl. 6, figs, 8-9. Hale. 1929; 280,
Figs. 1-4.
Cymodoce yaimardii, posterior region, Fig. 1.—Female (11 tm long) from Pearson T. Fig.
2 -Female (13.5 mm long) from Port Phillin (after Naylor), Fig. 3—Male (24 mm Jong)
from Western Porl. Fig. 4.—After Baker: "Prohably » young male"; locality and scale rot
Indleated,
CRUSTACEA; ISOPODA
fiz, 280. Nierstrasz, 1931, 205. Naylor,
1966; 189,
Naesa antennatliy
nomen nudum,
Cilicaea anrennaliy While Miers, 1884+
310,
Cilfcoaea = avitennalis = Miers,
1905: 335. Niersteasz, 1931: 205.
Localities; W. Aust.: Swan River (White
1847, Miers 1884) Vic.: Port Phillip
{Haswell 1881b. 1882; Naylor 1966); “very
common in shallow water around [southern
Australian] coasts" (Hale 1929). New
record, Pearson 1. (R at 25 m).
Material; Qne adult male (14 mm long).
Remarks: The specimen agrees with Baker's
(1908) and Hale’s (1929) descriptions, except
that the uropodal cxopods are ruunded
apically, rather than slightly bifid. No deserip-
tion tefers to the pads of short setue lining the
incurved inner surfuce of the distal part of
these cxopods and of the median projection.
This feature, together with the shape und
atrangement of these three projections, seems
to imply some definite function, such as clasp-
ing the female during mating, or clasping the
anterior region When the aninsal is rolled.
White, i847: 105;
Stebdbing,
Cilicaes Jatreillei Leach, 1818: 342. Miers,
1884: 308. Stebbing, 1905: 36, pl, 8,
Hiatle, 1929; 282, fiz. 282, Nierstrasz,
1931: 204. figs. 92-96, Naylor, 1966;
L90, fig. 3.
Neaesea latreiilei Milne Edwards, 1840:
218.
Cilicvea crassicandeta Haswell,
475, pl. 17. fig. 3-
Localities: There are numerous records from
South Africa, Ceylon, East Indios, Australia
and New Zealand [see Nierstrusz 193),
Naylor 1966). New recerd:; Pearson I. (R
at 20-25 m),
Material; Qne specimen, apparently female
and very young {6 mm),
Remarks: The females of Cymodoce pubescens
and Cilicaea latreitle’, both of which have bifid
twopodal exopods and are otherwise very simi-
lan. have caused much taxonomic confusion,
The Peurson I. specimen Jacks the characteria-
lic scale-sctae of Cymnoedoce pubescens, being
fairly liberally covered with stiff, erect setae:
it agrees well with Naylor's (1966) figure of
a female of Cilicaea latrejllei, although it has
not the well-defined anterior buss of the Port
Phillip (and Western Port) specimens, and the
posterior tip ef the uropodal exopod is forked’.
1881a;
abs)
These could well be juvenile features: com-
purison with a series of specimens from Wes-
tern Port leaves little doubt that it is a yery
young female of Cilicaea Jarreillei.
Jt must be noted thal the male figured by
Niesstrasz, despite his statement that “Die Tiere
(Figs, 92-96) stimmen put mit den Beschrei-
bungen von Miers [1884], Stebbing [1905], und
Barnard [1914] tiberein”, clearly belongs to
another species, and what he has labelled as
the female of C. latrei/le? is not a female of
that species, although it could he a Votng
male, Reliable figures will be found in the
papers by Barnard, Naylor and Stebbing; they
agree with all Victorian specimens available.
Genus CILICAEOPSIS Hansen, 1905
Hansen established this genus by designating
Cilleaca granulata Whitelegge (1902) as the
type, and his key to genera shows the diagnos-
tic characters to be; “Abdominal notch semi-
circular, without any vestige of mesial lobe.
ndp, of urp. rudimentary in the tale", This
seems to have been broudly interpreted as to
both the semicircular nature of the abdom-
inal notch and the rudimentary nature of the
endopod. Some of the species included in this
YeNUsS appear to necessitate a new generic diag-
nosis, but it will be best if modification of the
agnosis is left until this and the several
closely-related genera are reviewed.
Taking a broad view of the meaning of
semicircular, as Baker (1926) has done, the
new species described below conforms with
Hansen's diagnosis,
Cilicaeopsis flaccesa n,sp.
Locality: Pearson 1. (R at 25 ma: “From
algae on horizontal face”).
Material: One specimen, the holotype male
(median Jength 12,5 mm, total Jength 16
mm, width 6.1 min), wpparently adult, § A,
Shepherd, 10.i,1989 (NMY, J-249).
FIGS. 5-11
Diagnosis: Citicgeaopsis: with slender, curved
tropodal exopods beuring a Furry tuft of setae
on the median aspect of the distal end. Simi-
lur selae cover the dorsal surface of the endo-
pods and of the pleotelson below and behind
the two Jarge tuberculate bosses; surface else-
where glubrous. Appendix musculina Jong,
with a curved narrow tip extending well
beyond the setae of the second pleopod; inner
edge of the endopod grooved behind ta accom-
miodate the upper part of the appendix. Penes
long, tapering to a point. taterally compressed
and kinked backward near the end, Abdominal
W. F. SEED
Imm
FIGS, 5,6,7
5mm
FIGS. B. 9.
10, 11
Figs. 5-11. Cilicaeapsis floccosa. All figures from the holotype male. Fig. 5—Left maxillipede, Fig. 6.
—Left penis, from the right side, Fig. 7—Left second pleopod and appendix masculina.
Fig. 8.—Head, ventral view. Fig. 9. Whole animal, dorsal view. Fig. 10.—Posterior region,
from ihe left side. Fig. 11.—Pleotelson and uropods, ventral vicw.
CRUSTACEA: ISOPODA
notch wide and shallow, not visible from
above. Epistome pointed, much shorter than
broad.
Genus CYMODOPSIS Baker, 1926
Cymodopsis sp.
Lacalitys Pearson T, (R at 25 m, from algae
on horizontal face).
Material; One immature male (8 mm Jong),
with very rudimentary penes and appendix
masculina not yet appyrent-
Remarks: The specimen superficially resembles
Baker's (1926) figure of Cymodopsis crassa
in general form and in having two large coni-
¢al projections on the pleotelson. In lateral
yiew these are seen to be slightly undercut
below. forming a postcro-dorsal point on cach,
whereas the corresponding part of Cy. crassa
appears, in Baker's figure, to be smoothly
rounded and to run down to the telson as a
straight ftidge. Two other conspictious dif-
ferences are that the epistome is very much
shorter than broad (and different from that of
uny described species), and that the uropodal
exoped is relatively Jarge, plate-like, and
rejches to the end of the endopod in the ¢losed
position,
It seems clear that this is a new species, but
no sauistactory description can he given in the
absence of an adult male.
Group BURRANCHIATAE
Genus CERCEIS Milne Edwards, 1840
Cerceis aculicaudata (Haswell). Hansen, 1905:
127, Hale, 1929; 300. Nierstrasz, 1931:
214,
Sphaerama (9?) acuticaudata Haswell,
T8R1b; 197, pl 3, Fig. 9,
Localities: Vic. Griffith's Point, Port Phillip
(Haswell L&K1b); “This is a common
apectes” (Hale 1929), New record; Pearson
I. (R at 25 m, from algae on horizontal
face. R at 20-25 m).
Material: Three females (8.5-9.3 mm long),
all immature and Without ooslegites or
eggs.
Remarks: Specimens agree with descriptions
and figures of Cercels aeuticaydata except that
they lack the spines on the Urupods and the
spine on the pleotelson Is represented only by
1
a smooth median boss. Comparison with a
series of specimens from Western Port shows
that the growth of these spines is both allo-
metric and variuble.
Genus HASWELLIA Miers, 1884
Haswellia anomala (Haswell). Baker, (026:
273, pl, 48, figs, 8-9. Naylor, 1966; 192
Sphacroma (2?) anormnala Haswell, 188ba.
473, pl. 16, fig, 4,
Zuzara emarginata Haswell, 1881 b: 188,
pl. 3. fig. 5,
Taswellia emarginata (Haswell), Hansen
1905: 127. Hale, 1929: 304. fig. 304.
Localities: N_S.W.: Port Jackson (Haswell
188la): Vie.; Western Port (Haswell
L881b), Port Phillip (Naylor 1966); 8.
Aus! St. Vincent Gulf (Hale 1929). New
record: Pearson I. (R at 25 m).
Material; Seven females (5.88.6 mm long),
all immature and without eges, oostegites, or
modification of the mouthparts.
Remarks: The females of this. species are very
similar to those of Cerceis trispinosa. The uro-
poual exopods provide a convenient diagnostic
feature; in C. sispinasa females they are
longer than the endopods, while in A. anemate
they are both slightly shorter than the endo-
pods, and conspicuously toothed on the distal
edge. Comparison with a series of A. anomala
from Western Port confirms the identity of the
Pearson I. specimeus. The largest of them have
the hind. margin of the seventh thoracic tergite
produced in the centre (although not as fay as
in Haswell's figure of Sphacroma (7?) ana-
mala), a feature which supports Naylor's view
that S. (7?) anomala was the fernale of this
species and hence also supports his adoption of
the specific epithet anomala.
Haywellia cilicioides Baker, 1908: 158, pl. 10,
figs, (2-23. Hale, 1929: 304, fig. 305.
Localities: S, Aust; St. Vincent Gulf (Baker
1908, Hale 1929), New recerd: Pearson T-
(S at 30m).
Marerial: One adult male ( 9 mm long),
Remarks; The specimen agrecs with Baker's
(1908) and Hale's (1929) descriptions and
figures, except for slight differences in the
shape of the uropodal endopods, and in the
shape of the process of the Jast thoracic seg-
ment when viewed from above.
21>
Ww, F. SEED
Acknowledgements
Tam grateful to the Director of the Nationul
Muscum of Yictoria for facilities for this
study, und to the Science and Industry Endow-
ment Fund (C,S.1R,.0.) for a grant-in-aid.
Mrs. J. BE. Watson und Mr, A. Neboiss read
the draft manuscript and made a number of
helpful coniments.
References
Bagel, W. H, (1908), -Notes on some species of
the isopod family Sphaeromidae from the
South Ausiralian coast, Trans. R. Soc. 8.
Aust, 32, 138-162, pls. 3-10.
Baker, W. H, (1926).—Speciés of the isopod
family Sphaeromidae from eastern, southern
and western coasts of Australia. Trans. R,
Sac. S- Aust. 50, 247-279, pls. 38-53.
Bakrr, W. IT. (1928) —Australtan species of the
isopod family Sphaeromidae (continued).
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Barnarn, K. H. (19'4)—tConirihutions to the
Crustacean Fauna of South Affica, 3.—Addi-
tions. to the Marine Isopoda. with Notes on
some previously incompletely known Species.
Ann, S. Afr. Mus. 10, 32Sa—358a, 359-442.
pls, 27-38.
Hare. H. M, (1929).—"The Crustaceans of South
Australia”. Part 2, (Government Printer:
Adeluide: )
Hansen, H. J. (1905)—On. the Propagation,
Sirugture and Classification of the Family
Sphieromidae. Q. J] miarose, Sei, 49, 1s.
(193), 69-135, pl, 7,
HASwWeit, W. A. (1881a)—On some new Ans-
tralian Marine Jsopoda. Part I. Prac. Lite
Soe, NSW, 5, 470-481, pls. 16-19,
Haswrib, W. A. C€§88Ib)—Gn some new Aus-
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Hurry, D, E. (1961).—A Checklist and Key
to the Crustacea Isopoda of New Zealand
and the Subantarctic tslands. Trans. R_ Soe:
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Teach. W, LE. (t&13-14).—Crustacenlogy. Jn
“Rdinhurgh REncyclopyedia* 7. (Blackwood
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naire des Sciences Naturelles” 12. (Levrault;
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Munvins, R. J. (1962).—The Zoogeogranhy.
Ecology and Systematics of the Chilean
Murine Tsopods. Acra Univ. lind., NvF. Ava.
1, S7(11), 3-162.
Menzies, R. J... & PRANKENBERG. D. (1966).—
“Handhook on the Common Marine Isopod
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Miers, E, J. (1/884). -Crustaces. Jn “Report on
the Zoological Collections made in the Tndo-+
Pacific Ocean during the voyage of H.M,S,
‘AlerU, 1861-2," pp. 178-322. pls. {&-34.
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des Crustacés” 3. (Libraire Encyclopedique
de Boret: Paris.)
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1963. Isopoda, Afem. natn. Mus. View, 27,
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Watson, J. E. (1973)—Pearson Island Bxpedi-
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‘Thetis’. Crustacea Part TI, Tsopoda Part Il."
(Australian Museum, Sydney.)
STRATIGRAPHY, STRUCTURE AND METAMORPHISM OF THE
KANMANTOO GROUP (CAMBRIAN) IN ITS TYPE SECTION EAST OF
TUNKALILLA BEACH, SOUTH AUSTRALIA
BY B, DAILY* AND A, R, MILNES}
Summary
DAILY, B., & MILNES, A. R. (1973). -Stratigraphy, structure and metamorphism of the
Kanmantoo Group (Cambrian) in its type section east of Tunkalilla Beach, South Australia. Trans.
R. Soc. S. Aust. 97(3), 213-242, 31 August, 1973.
An apparently conformable sequence of metaclastics forming the upper part of the type Kanmantoo
Group (Cambrian) has been mapped along 50 km of coastline between Tunkalilla Beach and
Middleton Beach.
Sporadically occurring black carbonaceous and sulphide-rich phyllites are confined to and
characterise the Brown Hill Sub-group. Associated immature "flysch-like"” metasediments were
deposited very rapidly in an actively subsiding basin by currents flowing mainly from the NW.
Numerous pebble beds, similar to those in the underlying Inman Hill Sub-group, reflect the
continuity of the Kangarooian Movements. The cross-bedded Middleton Sandstone at the top of the
overlying Wattaberri Sub-group was deposited by currents directed mainly towards the east.
STRATIGRAPHY, STRUCTURE AND METAMORPHISM OF THE KANMANTOO
GROUP (CAMBRIAN) IN ITS TYPE SECTION EAST OF TUNKALILLA BEACH,
SOUTH AUSTRALIA
by B. Daity* and A. R. MILNES}
Summary
Datry, B., & Mines, A, R. (1973).—Stratigraphy, structure and metamorphism of the Kan-
mantoo Group (Cambrian) in its iype section east of Tunkalilla Beach, South Australia.
Trans. R, Soc. 8. Aust. 97(3), 213-242, 31 August, 1973.
An apparently conformable. sequence of metaclastics forming the upper part of the type
Kanmantoo Group (Cambrian) has been mapped alone 450 km of coastline between Tunka-
lila Beach and Middleton Beach.
Sporadically occurring black carbonaccous and sulphide-rich phyijlites are confined to and
characterise the Brown Hill Sub-group. Associated immature “fiysch-like” mictasedimenty were
deposited very rapidly in on actively subsiding basin by currents flowing mainly from the
NW. Numerous pebble beds, similar ta those in the underlying Inman Hill Sub-group, reflect
the continuity of the Kangarooian Movernents, The cross-bedded Middleton Sandstone at the
top of the overlymg Wattaberri Sub-group was deposited by currents directed mainly towards
the east.
Kanmantoo Group rocks younger than the Middlcton Sandstone are unknown in the Mt.
Lofty Ranges, Because the formation is intruded by the Encounter Bay Granites, a thick
cover must have been present at the time of intrusion and metamiurphism.
Warm casts were the only fossils located in the study area und are suggestive of a marine
environment of deposition.
The sequence mapped forms the eastern limb of a regional anticline overturned towards
the NW. Two phases of folding are recognised. F; folds plunge shallowly towards the SSW
er NE. Eastwards from ihe core of the highly appressed and asymmetrical regional anti-
cline, F; folds in metasandstones become progressively more open, symmetrical and upright.
Mesoscale Fo folds with E. to SH plunges are confined to the eastern patt of the type section.
A weak crenulation of S$; is observable near Coolawang Creek, and becomes progressively
stronger towards the east.
The Encounter Bay Granites were intruded and had crystallised prior to the main phase
of FP, deformation because thin granite sheets, concordant with bedding, developed the §;
and S» schistosities during the folding episodes,
Stability fields for the observed metamorphic mineral assemblages show that the meta-
morphism of the Kanmantoo Group was effected at moderate temperatures (below 540°C)
and Tow pressurcs (below 3 kb). This is indicated also by the well preserved sedimentary
sftuctures found throughout this sequence of undalusite-staurolite prade rocks, Cordierite,
restricted to the proximity of the granite, records the highest grade of metamorphism within
the type section. Petrographic evidence suggesis that cordierite crystallised during the pre-
to early syn-Fy phase, a conclusion in harmony with the suggested pre-tectonic emplacement
of the Encounter Bay Granites. Petrographic evidence indicates that most of the sndalusite
formed during the late syn-F; phase of metamorphic crystallisation. In the post-F, and
pre-F. static phase of meftamarphic crystallisation, some andalusite, garnet, hornblende,
*Centre for Precambrian Research, Department of Geology and Mineralogy, University of
Adelaide, Adelaide, S, Aust. 3001,
+ GS.1R.0. Division of Soils, Glen Osmond, 5. Aust, 5064, Formerly, Department of Geology and
Mineralogy, University of Adclaide.
214 B. DAILY and A. R. MILNES
scapolite and chlorite grew across the S; schistosily, The 8. crenulaiion cleavage, where
developed, deformed porphyroblasts of this age. Albitisation postdates the Sw schistosity.
A peculiar “striped” layering. post-S; and pre-S. in age, occurs within the Petrel Cove
Formation west of Rosetta Head and resulted from alteration adjacent te tensional fractures.
Two distinc! groups of basic dykes. post S; and pre-S» in age. cut Kanmantoo Group
metasediments.
Introduction
A previous paper (Daily & Milnes 1971a)
discussed the geology of the Jowcr part of the
‘ype sezlion of the Kanmantoa Group (Sprigg
& Campana 1953) as found in the coastal clitls
of Fleurieu Peninsula between Campbell Creek
and the western end of Tunkalilla Beach (Figs.
1 & 2). The present paper gives the results of
our investigations of the geology for the re+
mainder of the type section east to Rosetta
Head near Victor Harbor. In addition, we have
extended our averse to include stratigraphi-
cally younger metasediments to the NE in the
Port Elliot and Middleton areas.
The stratigraphic succession und geological
structure over this 50 km of coustline (Figs,
3 & 4). differ significantly from that shown
on the map given by Madigan (1925), on the
Jervis (Campana & Wilson 1954) and Encoun-
ter (Crawford & Thomson 1959) 1;63,360
Carrickelnaa Hes
Normanwilie
2
hy
Le]
555
VR
Canneshaw
Afwadle Arver
KANGAROO ISLAND
Fig. LL
Figs, 2-4,
map sheets, on the BARKER 1:250,000 map
sheet (Thomson & Horwitz 1962) and in the
‘Handbook of South Australian Geology’
(Thomson 7m Parkin 1969). In essence we have
found that progressively younger formations
occur to the east despite the advent of folds
which repeat parts of the sequence. These: folds
dre inclined to upright and plunge at relatively
low angles towards the SSW. hut as shown on
the maps, some exceptions in plunge direction
do occur, particularly in connection with parts
of the Wattaberri Sub-group.
Our main aims in studying the geology of
the type Kanmantoo Group have been to te-
assess. US stratigraphy and age limits and to
try and determine the relative time of emplace-
ment of the Encounter Bay Granites in con-
trast to the isotopic ages obtained by Dasch
ef al, (197), In the presumed absence of diag-
nostic foszils within the type section, we have
a
a
=
n
a)
uy
io
Houghton =
> ex
&
ADELAIDE a
ao e uneaa
Naifne e Brukungea
\ « Kanmantoo
I Call +45,
oc Sa, 2 Callington
tr Barber C4
K
Selick Hil A eo Wit Cornpass
cut Hil
ee a)
got alaman HiRes
«
ue Hindmarsh Valley
co
Ass,
“SA
DUDLEY’ FEMINSLULA,
~G Willoughby
© Hart
20 K'LOMETRES
bent el ee I
Locality mup of geographic features. exclusive of those shown on the geological maps in
GEOLOGY OF TYPE SECTION, KANMANTO0 GROUP (CAMBRIAN) ON
been forced to define the relative age of this
emplacement i tetms of the structural and
metamorphic history of the Kanmantoo Group,
Fortunately, sufficient rock samples were col-
lected and enough structural observations were
recorded during our traverse jo enable us to
siggest the emplacement of the granites prior
to the culmination of the first phase of defor-
mation.
We have not attempted a rigorous analysis of
the structure and metamorphism of the Kan-
manloo Group it tly Lype section, and further
work to this end is warranted, However, oite
of us (Milnes unpub.) has discussed “The En-
counter Bay Granites and their enviranment’ in
greece delail in a Ph.D, thesis, aubmitted to
the University of Adelaide.
Stratigraphy
In presenting the geology of the upper part
of the Kanmantoo Group we have revised the
stratigraphic terminology previously applied in
the type section (Daily & Milnes 19729). This
scheme is given in Table {, As explained in
the 1972 pauper, we have discarded the term
TABLE. |
Stratigraphic scheme for the Kanmantoa Group tn
ity mpe section between Campbell Creek anil
Rosetta Head. The table also includes strotigra-
phicaily younger metasediments ia the Port Eltiot-
Middteion arva and alsa shows the relationship
af the Kanmanioo Group ta-elder rocks (Precam-
brian and Cambilan) on Fleurieu Peninsula, The
regional unconformity between Precambrian and
Cambrian strata represents roughly the stratigra-
phie interval from the ABC Quartzite of the Flin-
ders Ranges to the hase of the iransgressive Carm-
brian sleposits-
Middiciwn Sandstone
Waraherel
Sub-group
Petre! Core Formation
Belauhidder Farmaion
Town Flu
Snb-eroun
Tunkalilla Formation
Tapanaiiya Farmition
KWanmanion Grain
Talisker Cale-siltsnone
Cambrian Sysivii\
Tamz2c Hill
Sub-aroun
Wacksiairs Passage Formation
Cavitkalinga
Alvad
Foviation
Campana Creek Mumher
Biiwhnte Crock Sillfione Member
Madigan Inlet Meniber
Normunyvilte Group
Remonal Unheonlinity
Marinn Group (Laie Preegnibrignl
Brukunga Formation but have retained the veo.
graphic terms Inman Hill and Brown Hill to
qualify sub-proup names, This procedyre re-
tains the position of the boundary between the
Taman Hill Formation and Brown Hill Beits
as desigaated initially by Forbes (1957), More-
over, the base of the Brown Hill Sub-group,
which is easily recognised, docs not correspond
to the base of the Nairne Pyrite Member at
the hase of the type Brukunga Formation. The
base of the Brown Hill Sub-group is estimated
In be 3000-4000 m above that position in the
Nairse region where the Brukunga Fornsation
was originally detined,
The following abbreviated gocount has been
made from our field notes of the coastal sec-
tions and environs east of Tunkalillu Beach
A. INMAN HILe Sup-croup (new rank)
The Inman Hill Sub-group (Table 1) con-
tains the Backsteirs Passage Formation,
Talisker Crilc-siltstone and Tupanappa Forma-
tion, All three formations occur in superposi-
tion in their type sections. In addition, we have
now located them on Dudley Peninsula, Kan-
guroo 1. (Daily & Milnes 1971b) where they
appear to be of identical facies. However, we
have had difficulties in identifying the Tapa-
nappa Formation in the northern sections of
Fleurieu Peninsula, although Tapanippa For-
mation of comparable lithology to that ex-
posed in the type section is well exposed along
the Mount Barker Creek, near Callington, fur-
ther to the north. So far we have not mapped
any intettongucing relationships between typical
Tapanappa Formation and other formations,
nor have we recognised fault relationships that
Might account for jis apparent absence in some
sections, Nevertheless, we believe that such
possibilities do occur and we have recently
undertaken investigations in key areas to solve
these problems,
Tapanappa Fermation
Detailed observations on the lower part of
the Kanmantoo Group (Daily & Milnes 19714),
ditt not extend beyond the western end of
Tunkalilla Beach where the modem coastal
clifis abut upon the beach, However. there ix
an arcuate line of old coustal cliffs behind an
alluvial bench backing that beach which per-
Mitted collection of data for the uppermost part
of the Tapanappa Forniatiun, despite the rela-
tively poor outereps in comparison with those
in the thodern coastal cliffline, The sequence
Cunsisty of dark coloured, thick-bedded to
laminated, generally fine- to coarse-grained
metusandstoncs, although near the top of the
Th
formation ver¥ fine-grained metasandstones
predominate. The individual metasundstone
beds, rarely more than 1 nm thick, are split by
povrly eutcropping and {frequently Juminated
octasiltslones and phyllite interbeds which are
penerully much thinner than (he metasand-
stones, These pelites range down tu partings of
i few millimetres. Poorly outcropping intervals.
many in excess of 10 m Stratgeaphic thickness,
are presumed metasiitstones or phyllite beds.
Bunds und tods ol cale-silicates occur spora-
divally in the metasandstones and even in some
of the finer grained lithologies. Porphyroblasts
oft chiurite. muscovite and bielite dre common
in the finer grained lithulogies, especially the
phyllites, ws well as some of the metasandstone
bands. Onternps of small-scale conglomerates
uceur ws thin discontinuous bands in same uf
the coarser grained metasandstones but their
presence in the sequence was generally inult-
cated by float rather dha outcrop,
B, Brown Hit. Sup-croue (new rank)
The base of the Brown Hill Sub-group and
the Tunkalill Pormatioi is marked by a
sequence of dark blue to black laminated phyl-
lites ybout 15 m thick (not 10 m as in Daily &
Milnes 197 1a, p, 207), and outcrops on a small
suddle hehind the beach about 2.5 km west
of Tunk Head. The Tunkalilla Formation to-
gether with the overlying Balquhidder Forma-
tion constitute the Brown Hill Sub-group. ‘The
lithologies within the sub-group ure not unlike
those found in the Tapanappa Formation and
indeed jt is impossible to: assign limited se-
quences to either unless the dark blue-black
carbonaccous and sulphide-rich phyllites cha-
racteristic of the Brown Hill Sub-group. are
located within them. Fortunately, dark
coloured phyllites of this type are unknown to
us guiside the sub-group within the type ares.
Geopraphicolly, they ure quite extensive as they
are known from Middle River (upstream from
the dam site} on Kanyguroo L, liorth to at least
ihe Callington areca in the eastern Mt. Lofty
Ranges.
Tunkalilla Formation (new name}
Vhe dark coloured laminated phyllites rark-
ing the base of this formation conformably
overlie the Tupanappa Formation metiasédi-
ments, When weathered, the phyllites are
characteristically stained yellow and brown by
jarosite ond guethite due to the oxidation of
iron sulphides within the Tock, Stratigraphically
above this basal unit are about 30 m of poorly
cutcropping dark coloured medium- to course-
grained mactusandistones with interbedded meta-
B. DAILY and A. R, MILNES
siltslunes und phyllites. Succeeding this there is
a khaki to mid-grey coloured phyllite about
30 m thick overlain by a 200 m thick sequence
of alternating Lo m thick bands of fine- to
mediuin-zrained melasandstones und Khakl to
mid-grey coloured phyllites. ‘These are in turn
overlain by a thick sequence (ahoul 150) a}
of predominantly well laminated metusiltstones
(Fig. 8) andl phyllites with minor metusand-
stone interbeds cut-and-fill into the finer clas-
tics, This interval vecupies: low ground and
outcrops poorly. Elongate to ovoid porphyro-
blasts wf micas iunlchlorite are Cummon in the
metasiltstones. Several small quartz-feldspar-
chlorite-muscovile pegmultiluy oocur in silky
phyllites near the mouth of Tunkalilla Creck.
A well developed crenulutvon is present in the
phyllites immediatcly adjacent to these ‘sweat’
Peymaliles, but this is local and has nov been
vbserved elsewhere.
The topmost part of the Formation is a well
laminated phyillite-metasilistone to fine-grained
metasandstone sequence capped by sc least 3 m
of blue-black luminsted carbanuccous phyllites
in which the alteration of sulphides. has. pra-
duced jurosite and poethite. Cubie shaped voids
after pyrite are readily upparent in the wea-
thered phyllites. The best outerops can be in-
spected high up on a mainly soil-covered west
facing cliff about 100 m east Of Tunkalilla
Creck, after which the formation is named.
The position of the upper boundary of the
Tunkalilla Formation is plotted on ihe geo-
Jogical map (Fig. 3). In the first creek west of
Cullawonga Creek. a & m thick band of black
sulphide-rich finely laminated phyllites stained
With jarosile and geethite marks the top of the
formation, It is underlain by about 3 m uf
metasinistone., Below the melasandstune there
is a much thicker poorly outcropping bund of
blue-black carbonaceous and — sulphide-rich
phyllites which, providing there are no struc-
(ural complicahions, may haye a thickness of
30 m. Paler coloured phyllites and metasilt-
stones occur below this. We may have missed
this approximately 30 m band of black phyllite
in the type section due lo thick soil gover at
this stratigraphic level. In craverses east of Cul-
lawonga Creek however, we Jocated only the
upper of the tWo bands and therefore i is more
Jikely that this thick band is a local develop-
ment.
The only fossils found within the formation
dre abundant worm casts in outcrops of sand-
blasted metasandstenes at the back of Bolla-
parudd’a Keach, adjacent ba the westeen hank
GEOLOGY DF TYPE SECTION, KANMANTOAO GROUP (CAMBRIAN) ZL
of that creek, The worm easts (Figs. 9 & 14))
are exceptionally well-preserved despite the tec
tonisia and metamorphism. ‘They are weathered
out in full relicE in an ourgrop about 8 m tong
and their tubular and sinuous nature can be
seen to pericction, particularly in sections nor-
mal to the bedding. The bioturhared interval,
Which is obour 20 cm thick, forms the topmost
bed in a large scour-channel (Fig. 14). The
lower parts of the channel comtain shale-chip
conglomerates in which the now phyilite frag-
mcals are scattered in a metasandstone matox.
Thus the Tunkolilla Formation in its type
section ut Tynkalilla Creck is a sequence (at
levtst 250 mM thick) of muinly fine-grained clas-
tics whose base and top are marked by bluc-
black carbonaceous and sulphide-bearing Jami-
nated phyllites of « characteristic appearance,
In the region to the narth of Hindmarsh Valley
mapped tty Forbes (1957), 4 seemingly iden-
tical sequence can be found on Mr, J. Grevn's
property just over | km SE of *Panybuly”
homestead (see Milang 1:43,000 map sheet,
Horwitz, & Thomson 1960, fo) the geographic
position of “Pambula''),
Balquhidder Formation (new name)
This formation is named afler the property
known as “Balquhidder (see Fig. 3), Its lype
section spans the coastline from a puint
approximately 2 km west of Tunk Head ease
te a point about 0.7 km west of King Poine,
where the much finer grained metaclastics
marking the base of the overlying Wattaberri
Sub-group are first encountered. There is con-
tinlOus Outcrop over this upproximately 30 km
of coastline except for the two stretches occu-
pied by Parsons Beach and Waitpinga Beach
However, the sequence is repeated by folding
across these two intervals so that our observa-
tions have covered most, if not all, stratigraphic
Jevels within the formation. The oldest part of
the formation occurs within a southerly-phing-
ing broad synclinal structure whose axis les
immediately west of Tunk Heath
A sequence of medium- to coarse-grained
metasandstones with metasiltstone interbeds
vecurs at the base of the formation, These are
conformable with the underlying Tunkalilla
Formation, About 300 m straligraphically
above the base of the formation and west of
Tunk Head. there is a 10 m thick sequence of
black carbonaceous and sulphide-rich phyllites,
Exposures are normally covered by sand on the
coast between this outcrop and the base of the
formation, but a seyvenge can be examined ine
land in the ancient coastal cliffs. The bulk of
~t
the rocks are thick-bedded, medium- to coarse-
gramed metasandstones with thin interbeds of
metasiltstones or phyllites, However, there ure
intervals Where the metusilistones (up to 3 im
thick) are the dominant lithology and are inter-
bedded with metasandstones up jo | m thick.
Some thin blue-black carbonaceous and pyritic
phyllites were also nuted, Porphyroblasts of
chlorite and micas occur in many phyllite inter-
beds as well as in some fine-grained metasand-
stones.
The basal part of the formation is best seen
it coustal exposures, for example in a traverse
from Tunk Head east to Bollaparuilda Beach.
In these superior outerops the sequence has a
different character, For instance, many of the
seemingly massive metasandstones are well
laminated and crass-bedding becomes nore
apparent. Other notuble features include phyl-
lite-chip conglomerates (up ta 0,5 m thick)
which are associated with coarse-grained to
zranule-rich metasandstones in cut-and-fill
structures, load casts and associated flame
siructures, and beds with climbing ripples
(Figs. 11-13). Cale-silicute rods and hands are
also apparent (Piys. 15 & 14),
Above the lowest blue-black carboouceots
and sulphide-rich phyllites within the Balyu-
hidder Formation on both limhs of the Tunk
Head synclinc, medium- ta yery coarse-grained
missive anu laminated mectasandstones are do-
minant (Figs 17-19). The beds are variable in
thicknesa and generally less than | m_ thick,
ancl where this is the case the sequence has
somewhar flagey appearance (Fig. 20). How-
ever, there are intervals where metasandstones
are Up to 3m thick. Nevertheless, the thickest
bed recorded was. a 15 m thick band of
medium- to coarse-grained metasandstene
occurring adjacent to a thin black carbonaceous
and sulphide-rich phyllite within the uppermost
beds of the syncline, Thin bands and lenses of
small-scale pebble conglomerates and phyllite
chip bands occur sporadically through the
sequence and are generally associated with the
coarser sand mteryals. Small-scale sedimentary
structures are most frequently found in the
metasillstones and laminated phyllitic intervals
(Fig 21). Cale-silicate bands ad rods are
again common.
The black ¢arbonaceous snd sulphide-rich
phyllites are best seen on the eastern limb of
the syncline just vast of Tunk Head with the
highest band visible in a gulch immediately
west of the tazor-edged ridge constituting that
head, Sulphides concenrewied into cross-culting
218 fh. DAILY and A, R, MILNES
veins within this wnit at thiy focality were pro-
byhly remobilized at or subsequent to the ume
of metamorphism (Vig. 22).
Between Calluwonga and Bollaparudda
Beuches, the hagal part of the formatiun ts simi-
lar tw that described above, except that small-
scule Conglumetutes are present in some of the
cut-and-fill structures within thick mietasand-
stones. Phyllite-chip conglomerates are again
conimun ussaciales.
On the point 0.4 km SE of Bollaparudda
Beach, there are massive and well-bedded
course-grained metasandstanes, 15 em to 7 om
in thickness. ulternating with phyllite and meta-
siltstoie interheds. Many sands are bioturbated.
Higher in the sequence pebble bands (includ-
ing gaciss pebbles up to S cm across) and
lenses of small-scale conglomerates occur with-
in laminated medium- to coarse-grained: meta-
sandswnes (Fig, 231. Phyllite-chip congle-
merutes ure aussaciuted with these canglomer-
ates venerally near the bottom of chunnoels.
Additional pebble bands in very massive lonk-
ing mielasanustones occur higher im the 3e-
quence. At first sight some thick bands appear
lo be structureless graded units with pebbles
near the base and. with grain size fining up-
wards, However, in all cases the sands were
found in detail to be well bedded (Fig. 24).
The high incidence of conglomerates in the
lower purt of the formation cast of Bolla-
purudda Beach contrasts with their relative
paucity further west. The conglomerates arc
comparable with those found in the Tapanappa
Formation. Similar conglomerates in’ rocks
which we regard ay being well down in the
Balquhidder Formatiun occur in the Cut Hill
roud-eutting: on the Mt. Compuss-Victor Har-
bor toad. ‘Therefore, it seems thal conglome-
rates in this general stratigraphic position may
be a widespread feature of the formation, thus
refiecting the continuity of the Kangarooian
Movements (Duily & Forbes 1969: Daily &
Milnes 1971a) which were partly responsible
tor the deposition of the Kanmantog Group.
A byck carbonaceous and sulphide-rich
phyllite occurs near the cape about 1 km cist
of Bollaparudda Beach and is overlain by a
seyuience of anrinated to poorly-bedded meti-
sandstone beds {up to | om thick) with thin
phyllite and metasiltsione interbeds. From here
to the mouth of Coolawong Creck the strike is
almost parallel to the coast. A 3 m thick band
of black carbonaccous and sulphide-rich phyl-
fite overlain by thick poorly-hedded metasand-
stones and thin {nterbeds ol metasiltstanes and
khaki-coloured phylites occurs at the moulll of
Coolawany Creck This may be the same sel.
phide band 4s seen on the cape alluded to
above.
Between Covlywang Creek and Pitsons
Beach, a number of fuld hinges and minar
sults were loculed within the successiun. Mak-
ing due allawance for these, the sequence 's
seen to consist mainly of orassive Lo yell-
laminated metasandstones in the lower part of
the succession. Many of the channels cut within
the sands are filled with coulse-graincd to
granule-sized claslics including, Jenses ol sill
pebbles, Nodules aiid irregular shaped bands of
chle-silicate lithology are developed in some
parts of the sequence. About T kim east of
Coolawang Creck, porphyroblusts of scapolite
occur in slumped and laminated metasand-
stones and phyllites, [a some of the metasill-
stones hoth scupolite and garnet muy be found
In ihe sume arca pexmatites and feldspathived
vones are common and are offen associated
with tensional Features developed in) massive
metasandstone bedy split by thin phyllites ( Figs.
35 & 26). A litle higher in the sequence some
of the mussive metagandstones (1-2 m thick
sod split by phyllites up to 1 m thick) are
biolurhuted towards their tops. A weak créenu-
lation cleavage, noticed for the first lime in our
travetse, becomes much more apparent as one
moves to the east. und provides evidence for a
second phase of folding, The first-generation
folds are open and asymmetric with steep
westerly-dipping unticlinal limbs in contrast to
the inclined folds further west possessing over-
turned easterly-dipping anticlinal limbs, The
opening up of the fold hinges might he con-
sidered a function of distance wway from the
crystalline basement at the time of folding.
lhe remainder of the sequence vast to Par-
sons Beach is made up of flaggy laminated
metusandstones, generally less. than 1 m thick,
with interbedded thin laminuled and sumetimes
rippled metasiltsiones (Fig. 27) and ercnulated
phyllites up fo 0,6 m thick. Some of the sanils
are biowrbated und some contain cule-silicate
nodules, A feature of this pant of the formation
is the Fashion in which the cleavage has des-
troyed or vastly modilied the fine sedimentary
structures within the top (and bottom) few
centimetres of the sandier beds. Bedding sur-
faces, <imilar to thi shown in Fig. 19, show an
obvious lineation along the cleayage/ bedding
intersection duc to the smearing out of the sedi-
mentary siructures, Another expressiun of the
cleavaye/ bedding intersection 7s the pro-
GROLOGY GF TYPE SECTION, KANMANTOO GROUP (CAMBRIAN)
nounced ridge-and-furrow Tinention seen on
the base of some metasandstones (Fig, 28)
This simulates the cast of 4 glaciated pavement
ur sedimentary gfoove casts.
‘Yhe readily accessible sequence farming the
headland hetween Pyrsums Beaeh and Wait-
pings Beach is on the castern limb of a SW
plunging syncline occupied by Parsons Beach.
Massive und laminated coarse- to fine-grained
metasandstones with interbedded metusiltstones
und phyllites again constitute the succession.
In general, the metayandstone bunds increase in
thickness up the sequenee and near Parsons
Beach some beds pre nearly 6 m thick. Sedi-
mentary structures such as cross-bedding
faceasional sats up to 0,65 m), plunar-hedding,
slumping, convolute-bedding (Fig, 29), lond
gasts and jssociated Name siriictures, current
ripples, simple and compasite cut-and-fill struc-
tures {these are sometimes associated with thin
small-scale conglomerates und phyllite-chip
conglomerates), and bioturbated sandy inler-
vals were recorded. There wre several thick
phyllite and metasiltstone intervals. particularly
towards the bottom of the exposed sequence.
Some of these phyllites are crenulated and con-
tain cordierite and other porphyroablasts. Cale-
silicate bands. and rods, some of which are at
least 2 m long, are prominent through many of
the metasandstone intervals, Segregations pos-
sessing the sume mineralogy and encompassing
parts of pebble beds were also recorded.
There is a total absence of Kanmantoo
Group rocks across the 4 km wide sanily
Waitpinga Beach, Cainozeic und Permian
depusits (uncoliuured in Figs. 3 & 4) blanket
these rocks for many kilometres inland. How-
ever, the seyuence is continued uninterruptedly
along the coastline from just west of Newlund
Head to the vicinity of Rosetta Head.
On the western side of Newland Head there
ure very thick (up to 10m) medium- 19 coarse-
ernined metasandstones with pebble bands
which are mainly assuciated with out-and-fill
structures. The sands are well laminated and
cross-bedded within and away from the channel
hottams (Fig. 3M). Pebbles are scattered
through some of the hedded metasindstones
ind in One instance were seen to be present a>
the stoss side but absent on the lee side of
mego-ripples. A band of ceenulated andalusite
schist was found about 20 m east Trom the
commencement of oimcrop. Above this and
below a 2 mihick black carhonaceous anid sul-
phide-rich phvilife is a thick metasandstonc:
219
Stratigraphically higher ts a very thick sequence
of metasandstones (beds generally Jess than
2 m thick) which alternate with phyllites or
Thetysiltstones. ang an ocexsional much thicker
Metusandstone interval (Fig. 31), Two addi-
tioagl Black carbonaceous and sulphide-bearing
phyllites and two bands of andalusile schists
were located in this latter sequence, Strati-
stiuphically above and perhaps 300 m NE of
Newland Ueud, another prominent blue-black
curbonuceous and sulphide-rich phyllite is inter
bedded in the flaggy sequence (Fig. 32). This
can be readily followed for about J km before
it strikes jwland and is Jost from view. IL is
overlain by coarse metasandstones containing
weak developments of small-scale couglomer-
ales in an otherwise flaggy sequence, From this
point onwards to within about 2 km west of
King Point (Fig. 32), the strike of the beds
parallels or is slightly oblique to the precipitous
coastline, parts of which ure totally inacces-
sible. Consequently, our traverse was made
mainly along the tep of the coustal cliffs. An-
other slightly younger blue-black phyilite
occurs 2.4 km NE of Newland Hew and below
well-bedded, coarse-grained. impure metasand-
stones. split hy thin phyllites.
Aboul 1,5 kin WSW of King Beach a 2m
thick band of a comparable hlie-hlack sul-
phide-rich phyllite uceurs above a regnlarty
bedded medium- to coarse-zrained metasand-
stone with thick laminated phyllite partings, Tt
is either stratigtaphically above or possibly on
the same horizon as the blue-black phyllite
mentioned shove. Towards King Point lami-
nated metasandstones, often with cut-and-fill
structures, occur in beds varying from abeut
19 cm to § m in thickness. Banded metsilt-
stones aud finely crenulated phyllites (maxi-
mum thickness about 2 m1) occur as interberls
and may contain porphyroblasis of mica or
more rarely andalusite.
The uppermost part of the formation is
marked hy a sequence of relatively fiagay
metasandstones and tactasiltstumes, in which
there are a number of interbeds of knotted
undulusite schists, Cordicrite parphyrnblasts
andl =o quartz-tich uyyregites co-exist. with
undulusite in some of the schists (Fra, 347, ‘the
cordicrite porptyroblasts and quact¢-rich agere-
gales ute deformed in the plane o£ the pro-
minent mica schistosity (S,) resulting in the
development of small augen. Tn many pelitic
units a prominent discontinuous layering occurs
parallel to S, defined by the alignment of these
augen. Andulusile porphyroblasts. which are
22)
seen in thin section to postdate the augen
development, also help to define this layering.
Andalosite-rich pegmatites containing mus-
rovite and corundum occur within this interval,
Small-scale first- and second-generalion folds
ate commonly present m the metasediments
vdjioent to the pezmiatites, which are invariably
strongly boudinaged,
C WATTABERRI SuB-GROUP (new name)
A marked change in sedimentation took
pluce ufter the ueposition ef ihe Balquhidder
Fornilion duc to the influx of fine-grained
ctustics. These now constitute a sequence of
essenlially phyllites or schists, metasiltstones
und fine-grained metasandstones and are dis-
cussed below under the name of Petrel Cove
Formation, The succeeding characteristically
faminuted and cross-hedded Middleton Sanu.
Stone is Yrouped with the Petrel Cove Forma-
lion 45 the Wattaberri Sub-group. named from
the property called “Wattabers”. some 2.5 km
northeaf Port Elliot.
Perrel Cove Formation (new name)
This formution is much less resistant to ero-
sion than the Balquhidder Formation and thus
forms | more subdued topography, [t occurs
in low ¢lill lines between a point 0.75 km west
of King Beach and Petrel Cove. aftee which
the formation is named. Good outcrops can
also be inspected at low tide in Rosetta Huar-
boron the NW side of Rosetta Head. All parts
of the formation in this area are readily
accessible. It can be regarded as one of the
key areus in the State where so many aspects
of the effects of metamorphism and tectonism
of a sedimentary sequence can be demunstrated
clearly. All efforts should be made to preserve
this stretch of coastline as a geological monu-
Hent. Vanous aspects of the geotory of parts
of the sequence have already been discussed by
Browne (1920). Bowes (1954), Hobbs & Tal-
bor (14956) and Talbot & Hobbs (1968 and
TV69],
A sequence of folded porphyroblastic and
sindalusite-cordierite schists constitutes the hase
of the Petrel Cove Formation. These confor-
mably overlie the topmost unit of the Balqu-
hidder Formution which 1s composed of a
sequence of fine-grained metisandstones and
mejasiltytones, The schists exhibit a well-
developed gugen layering (S,) axial plane to
F, tolds. which are cleatly outlined by the
hedding (S\,) ws is seen in Figs. 35 and 36, Tn
the saihe Oulerops (ste especially Fig, 35) verv
thin light-coloured layers pervade the schists
and appea as a senes of thin closely spaced
B. DAILY and A. R. MILNES
stripes (S,) at a low ongle to the auger layer-
ing. These stripes define what is herein termed
a “striped” layering (see also Talbot & Hehbs,
1968), We regurd this “striped” layering as
having developed by alteration along tensional
(ractures post-dating the F, folds. Note also
that tn the same outcrop (Fig, 35) there arc
“pincth-and-swell” pegmatitic layers parallel io
the “striped” Jaycring and along which disloca-
tion hus taken place, In several areas more than
one set of “striped” layering can be recognised.
A well laminated metasilistone sequence
(Fig. 37). parts of which are strongly folded,
oecurs on the wave-cut platform pbove the
basal anulalusite-cordierite schists, Mics porphy-
roblasts, light-coloured augen m S,, and an in-
frequent “striped” layering show up in parts
of this succession. Dislocation of bedding paral-
lel to the stripes is ugain evident in some aul-
craps.. Stratigraphically above are porphyro-
blastic metasiltstones and schists with intervuls
Showing either isolated augen in S, or with
dugen sufficiently concentrated to produce an
augen layering in S;. Very fine-grained meta-
sandstones are interbedded with these. On King
Point the beds are well laminated and exhibit
small-scale sedimentary structures (Fig, 38).
Near this locality some calc-silicate layers occur
parallel to the bedding and shaw « cleavage/
bedding intersection plunging towards the
south, These beds strike across the bay rowards
Petre] Cove,
Along the coast between King Beach and the
contact hetween the Petrel Cove Formation
with the Encounter Bay Granites on Rosetta
Head, the incompetent metasediments within
the Petrel Cove Formation show numerous
meso-scule Folds of two generations, The folded
sequence consists of very fine grained meta-
sanistones with interbeds of metasiltstones and
crenulaled schists. Numerous deformed sepli-
mentary structures such us those in Pigs, 39
and 40 are common. Similar sedimentary struc-
tures have been figured or discussed by Hobbs
& Talbot (1966) and Talbot & Hobbs (1969),
The common occurrence of “striped” Llayer-
ing (Figs. 39-47) which, in a thick homo-
geneous rack type, could passibly be mistaken
for bedding has heen mentioned above. This
layering is markedly refracted across the
boundaries between the metisandstones and the
andalusite-cordierité schists. The layering is not
only discontinuous hut may even show a
feathering effect fPig. 454, aguin u feature sug-
gesting that rt developed in response to ten-
sion. In addition, it formed afler the F, feld-
GEOLOGY OF TYPE SECTION, KANMANTOO GROEP ICAMARIAN) 32]
ing phuse because faint stripes cut straight
through strictures on the tap of heds deformed
by that phase of folding (Fig. 39). In many
andalusite-cordierite schist intervuls, especinlly
along the coastline immediately west of Petrel
Cove, the “striped” lavering has been folded
on a amall scale about an axial plane purallel
10 the crenululion cleavage (S.) (Figs, 43 and
461. Many of these andalusite-cordicrite schists
show either a preferred concentralion ur a
depletion of andalusite porphyroblasts paraile}
to the “striped” layering (Fig. 47),
Cummozoic and Permian sediments (un-
colonred in Fig, 4) blanket mast of the region
between Rosetta Head and the hills lying to the
nerth of Port Elliot. Over this distance no rocks
helonging to the Wattaberri Sub-group are
known to Getcrop on the coast, However, in a
section between Brown Hill and Wattaberri
we have inferred a houndary heltween the Bal-
qguhidder Formation (containing some blue-
black carbonaceous and pyritic phyllites) and
an overlying dominantly phyllitic-metasiltstone
sequence identified as the Petrel Cove Farma-
tion (Fig. 48). Despile a prolonged search in
the area, we have been unable to locate anda-
lusite or other distinctive melamornphic
minerals so characteristic of many parts of the
formation in its type section between King
Tomt and Rosetta Head. Stratigraphically
above these racks are metasandstones of w dis-
unetive facres which we call the Middleton
Sandstone. This formation is best examined in
the Middleton quarry and on Middicton Beach
Metasandstones of an identical facies and
stratigraphic position lie in contact “ith the
Encounter Bay Granites on the eastern end of
Kangarcur 1,
We have been unable to locate a contact
between the Petrel Cove Formation and the
Middleton Sandstone une ta imperfect qutcrop
Nevertheless, 9 tentative houndary has been
positioned as shown in Fig, 4. Rocks shown as
Middleton Sandstone pre characterised hy the
presence of epidote-rich bands.
Middleton Sanestane (new name}
The best section of Middleton Sandstone is
seen on the wave-eut platform in the vicinity
of Middleton (Pig. 4). These outcrops. which
are totally isolated from other Kunmantoo
Group rocks, consist largely of grey-coloured
fine-prained metasandstones which are gener-
wily Ughter in colour and mure quartzese than
the bulk of the metasandstones found in the
Kinmunton Group, with ie exception of some
of the Inmun Hill Sub-group metasandstones,,
for example those of the Backstairs Passage
Formation, The metasandstones ure typically
very well-laminated and are commonly cross-
bedded wilh sets up to 1.2 m thick (Fig. 49)
and with an indicated current direction gener-
ally from the west throughout much of the
sequence. In some of the higher parts of the
outcropping sequence, more random current
directions are indicated, Some slumping down
the direction uf the cross-hedding is evident.
Breaking the monotony of thie well-laminated
sequence ure intervals with minor sedimentary
structures. mainly small-scale current ripples,
which include some starved ripples involving
material of somewhat different grain size.
There wre also rare metasandstone feds up lo
0.5 m thick containing angular pbyllitice clasts
up ta 30 em long and 15 cm thick, though
generally the clasts are platy and less than 5 cm
acress, They represent the ripping up of partly-
indurated mudstones by strong currents and
their subsequent depositinn in scour-channels
after very limited twansport. The only other
metasediments present are a Few metasiltstones,
up wo 10 m thick, containing chlorite and
actinolite,
One of the mast conspicuous features of the
sequence is the prevalence of pale-green
epidote-rich sepreeations which are developed
consistently throughout the succession,
Albitised bleached zones of alteration ate also
presence. Most of the epidote-tich segregalions
vecur eilher in bands or as lenticular patches
and nodules parallel to the bedding (Figs. 50-
S52) but some epidole occurs in association with
quartz-chlovite-feldspar pegmatites that are de-
veloped in fractures and boudinaged zanes
All beds in the sequence fuce north. Obser-
witions show that the frequent occurrence of
overturned southerly dipping beds is due solely
jo warping locally developed along the sirike.
Practute-cleavage/ bedding intersections indi-
cute that the heds belong to the southern lind
of a synelinc plunging shallowly towards the
east, This syneline is a second-generation fold
as Shown by overprinting relationships in
Midd'eton Sandstone in the Middleton qitirry
The recognition of Jarge second-generation
folds in this area significantly adds fo the small
number of macroscopic F., folds reparted fror
elyewhere within the Mr. Lofty Ranges hy
Offler & Fleming (1968),
No upper boundary to the Middleton Sand
stone is known on the Fleurieu Peninsula, and
consequently this formation comprises the
youngest Kanmanino Group rocks in the
322 B. DAILY and A. R. MILNES
eastern Mt. Lofty Ranges. Because the En-
counter Bay Granites exposed at Port Elliot
seem to have intruded rocks of this formation,
a substantial sedimentary cover must have
existed above the Middleton Sandstone at the
tine of granite emplacement and metamar-
phism. This cover is likely to have embraced
rocks ef Middle Cambrian to possibly Karly
Ordovician age, as rocks spanning this sugaes-
teil lime interval occur on Yorke Peoinsula
(Datly 1969) and in the Flinders Ranges
(Daily & Forhes 1969).
D. INTRUSIVE Basic Dykes
Two groups of basic dykes that have in-
(ruded the Kanmuntoo Group within the type
section wre shown in Figs, 2-4, The dykes of
the first group are transgressive fine- to
meditim-grained metadolerites, Similar rocks
have intruded the Encounter Bay Granites at
Rosetta Head and Port Elliot.
‘The second group consists of hasic dykes
exlensively contaminated by meta-sedimentary
rock material. One such dyke containing large
feldspar megacrysts. intrudes Balquhidder For-
mation metaxediments along the coast cast of
Tunk Head (Fig. 3). This dyke was described
by Madigan (1925), A second contaminated
husie dyke-rock occurs in the line of ancient
coustul cliffs backing Tunkatilla Beach, but it
is poorly exposed. Sirnilar contaminated basic
dykes have intruded the Middicton Sandstone
dong the south cous: of Dudley Peninsula
(Kingarco I), west of Cape Hart. The results
of our investigations of (hese rack lypes will be
presented tna subsequent communication.
Structure
A lar Structruse of He Mera-sepimMen-
TARY SEQUENCE
An Interpretation of the geological structure
of the Lower Cambrian rocks exposed alone
the south coast of Fleuricu Peninsula between
Campbell Creek and ‘Tunkallly Beach has been
even in Daily & Milnes (1971a). This
sequence. involving the upper parts of the Nor.
manville Group’ and the lower parts of the
Kanmantog Group, is contained within a NE
plunging regional anticlinal structure which is
overturned io the NW [Fig. 2}. On the normal
eastern limb of this fold between Madigan fn-
let aud Tunkalilla Beach, all the mapped
mucroseale folds are believed to be first-
generation (F,) structures, Generally they
conform to the style of the regional fold, and
indeed to the style of folds developed in the
Late Precambrian ‘lorrens Group rocks in the
Houghton area described as B, folds by Talbot
(1964), and most of the folds described by
Omer & Fleming (1968) as F, folds within
large areas of the Mt. Lofty Ranges. They are
mainly inclined and asymmetric with eastern
bmbs of unticlines longer than western lintbs.
and with axial planc cleavages which dip
steeply to the SE. In many instances, disloca-
tion of the overturned western limbs of -unti-
clines has taken place along faults thar tend to
purullel Lhe axial plane schistosity. All struc-
tural data for this section of coastline are sum-
marized in Fig, 5 which shows the following:
(9) poles to bedding (S,)) indicate a fold usis
estimated to: plunge towards 047° at a low
angle, although mesoscale F, folds plunge
al Variuble angles towards both the NE
and the SW:
(b) intersections of §, with cleavage (8,),
long axes (L,) of phosphatic nodules Cin
the Heatherdale Shale), and the elonga-
tio of boudins resulting from: the defor-
mation of cale-silicate bands within the
overlying Kanmyntoo Croup are parallel
ta the axes of mesoscale F; folds;
(c) F, fold axes and the long axes of phos-
phate nodules und ealc-silicate boudins
define a great-circle distribution which
may be the result either of refolding or of
inhomogencous strain;
{di a lineation defined by the elongation of
calcite and mica crystals on $, surtaces
{not heddiny surfaces as reported in
Daily & Milnes 1971u, p. 207) in eal-
eureous and pelitic intervals respectively,
pitches up to 20" more steeply than 1,,,
und is now interpreted as a first-gencra-
tion structure Li’ (not a-sccond-generation
xtruciure as suggested by Daily & Milnes
197ta). The preferred orientution of horn-
blende poikiloblasts and. chlorite porphy-
roblasts. on §, surfuces in cale-siliegte
hourtins. and pelitic tntervals respectively
is approximately parallel to Li’, but its
significunce is not understood.
' This new name is defined herein to inchide all the Lower Cambrian sediments between the bise
of the Mount Ternble Formation and the base ot the Carritkalinga. Head Formation as mapped
in the Sellivk Llill-Normanville area by Abele & MoGowran (1959) and revised by Daily (1963),
GEOLOGY OF TYPE SECTION, KANMANTOO GROUP (CAMBRIAN)
FOLD. PHASE PLANAR ELEMENT LINEAR ELEMENT
: S + surface
Spe peaning
Tihat generatien
elructures
+y S)=Schielmeity Ly 7 Uheation ar
eis ut dott iw
My with Sy as
welal su fface
by = olnerar
1 Plonyation in Sy
Second generation
alruciures
hy S.=—crenulalion
7 cleavage,
Asia! Burtace
ta Fy tots.
Lo bineation gr
Beis af told or
granulation ia,
or Wilh S;
ay ania Surtace™
[i renresents ¢elimated direction and plunge of fold aais,
wr
ah.
a Peo AyHO47 s1meeD®
Pa ms
A
Miage i" |
*., /
\ i 4. : - f
“\ t d y
~s
e Poles to Se
A Poles to S-
se ee + F Fold gees
2 &, Sleavage/hedding jntersecvons
VY lLoeasleite, cise slangailyds im Sy
Fig. 5. (Above)—Table of structural elements
and corresponding symbols utilised in the
text.
(Below }—Equal area projection of struc-
lural daty for the section between Madi-
gan Inlet and Tunkalilla Beach.
Structural data for the remainder of the type
section and the region east to Middleton are
as follows:
(i) Turkalilla Beach to Rosetta Head
All the macroscale folds mapped and most
of the mesoscale folus observed to the east of
Tunkalilla Beach are interpreted as F, struc-
lures, They cojiltrast with the style of the
regional fold ta the west (mentioned above)
in that the western anticlinal limbs are not over-
turned, The folds are however mainly asvm-
metric with western antichnal limbs stecper
and shorter than the eastern limbs although
towards Rosetta Head the folds tend to be
more upright, symmetrical and apen (Fig. 36),
In yeneral, folds in metasandstones are oper
223
structures Whereas folds in the metasiltstones
and schists. particularly within the Petrel Cove
Formation, are smaller scale structures that
tend to be moderately tight and upright. The
transition [rom one style to another within the
metasandstones cannot be pin-pointed, Our ob-
servations suggest that it is gradual, and it
seems significant that the highly appressed F,
folds are characteristic of the oldest parts of
the stratigraphic sequence, whereas folds in
progressively younger formations are more
open and upright. ‘Vhe validity of this proposi-
tion is strengthened becatise on Dudley Penin-
sula, Kangaroo L.,.and along the Mount Barker
Creek. comparable situations pertain on the
eastern limb of the regional anticline.
The axial plane schistosity in F, folds is
defined by the preferred orientation of micas
which commonly enwrap small quartz-rich
aggregates and, in some localities east of New-
land Head, cordierite porphyroblasts. The
resulting augen define a unique augen-layering
parallel to S, (Figs. 34 & 35; Talbot & Hobbs
b9G8). In the schists est of Newland Head
there is a distinct but often discontinuous linea-
tion due to the elongation of augen on §, sur-
faces (Fig. 53). ‘This lineation is decidedly
different in orientation from that of the inter-
section of S, and 8, surfaces and the axes of
F, folds.
A weak crenulation of S, is apparent in
phyllites near Coolawang Creek and bevomes
more distinct eastwards, This crenulution pro-
vides evidence for the overprinting of first-
generation structures, In some areas there is a
penetrative strain-slip cleavage (Sa) which is
axial plane to mesoscale second-gencration
(F.) folds, Such folds are most cofispicuous
adjacent to pre-F., metamorphic pegmatites
(Fig. 54), They tefold small-scale F, folds
(Figs. 54 & 55) and are responsible for the
local variation in the plunge of F, fold axes
from NE to SW. It should he noted that in the
Rosetta Head region, Talbot & Hobbs (1968,
p. 584) have pointed out “that several sets of
crenulation cleavage are developed lacally”, We
have mapped only one crenulation cleavage
(S,) in that area,
The distinctive “striped” layering (S,)
referred 10 above is best developed just west
of Petrel Cove, Talbot & Hobbs (1968) also
reported ifs occurrence in the Kanmantoo Mine
area in metasediments assigned by Duily &
Milnes, (1972a) to the Tapanappa Formation.
The “striped” liyers are zones of alteration and
consist predominantly of quartz and plagioclase
224
wilh seme muscovite. They are up to 3 tm
wide, anu ure often bordered by thin biotite-
rich zones. I tiddition they may show a median
biotite nu/ or qual'tz-filled fracture (Fig. 56)
Althaugh ihe “striped” layering cuts across S,.
i fellct Sy schistasity. outlined by the preferred
orientation of muscovite and rare biotite, can
be seen within the “striped” layering. Struc-
tural relationships show thar the layering ts
not only post, but clearly pre-Fa (Figs. 43
& 46), Talbot & Hobbs (195, p. 585) sug-
pested thal the “staped” layers are “hon dilata-
tional and represent same form of differentia-
tion process. in situ’. In wew of the presence
of a central fracture within the “striped” layers,
the refraction of Inyenng across bedding, and
Fouthering, we can only conclude that these
structures ure dilatational anid wre due to ten-
sion and subsequent alteration adjacent to the
Fractures,
Alt structural data. collected between Tunka-
lilla Reach and Rosetta Head ate given in Fig.
® a=d. This shaws the following: Poles 10 3S,
show a great-circle spread and indicate a [old
axis estimated to plunge at 25> towards 200°
(Fig. 6a), This is supported by the attitudes of
Ly lineations defined mainly by the intersec-
fiun of S, ad.S, surfaces (Fig. 6b), However,
uxes Of mesoscale Py folds are distributed
about Wo point maxima, indicating plunges
towards the NE and the SW at shallow angles
(Fig. 6c). This suriition in plunge together
with the significant spread of poles to S, (Fig.
fe) may indicate refolding of the first-genera-
ijon ytructures, or may be dug to differential
strain. Ls lineations (the axes of crenulations
jn S,) plot as a diffuse maximum giving an
estinvaled plunge towards 165" at 63° (Fig,
ful =
(ii! The Brawn Hill drea
In the Brown Hill area, large scule NE plung-
ing asymmetric F, folds in Balquhidder Forma-
Gun metisedimenis occupy 4 narrow NE trend-
ing zone. The folds gre upright, and have the
open style typical of F, folds in metasandstones
in the eastern part of the type section. The
axial plume schistosity is defined by the pre-
ferred orientation of micas, A lineation formed
py the intersection ol 8, and S,, surfaces is
parallel io F, fold axes, $, surfaces in pelites
contain a mineral lineation defined by the pre-
ferred orientation of micus. This lineation has
a sleep southerly pitch in $,, and is interpreted
B, DALLY and A. Ro MILNES
usu firsi-veneration seructure (Loy. It is best
seen in the Lincoln Park quarry, south of
Brawn Hill.
The structural elements meastired in this arca
ure plottel in Fig. Ge, Poles to Sy lie along a
greal circle, and imuicule an Fy fold axis esti-
muted to plunge at 35° towards U51°. This is
supported by the attitudes of 1,’ lineations.
Two measuted L,’ lineations plinge steeply
towards the south,
Gil The Middletant A rect
Fust of the Brown Hill area, Fy, folds in
metasediments assigned to the Petrel Cove For-
mation and the Middleton Sandstone have been
overprinted by large- and small-scale Hy folds
that plunge at shallow angles towurds the SE
The relationships between F, und Fy siruc-
lures, both of which exhibit varving deerves. of
development in the area, wre besr examined tn
the Micdkdlelton quarry wbout L4 km NW ol
Middleton.
SE plunging F.. lolds are the deminant siruc-
tures in the Middleton quarry, und are
moderately uppresstd, inclined folds im which
the northern limbs of anticlines are longer thin
the southern limbs, and tend to he overturned
(Fig. 57). The larger folds contain small-scale
refolded Fy folds in pelitic units (Fig, 48),
A prominent set of fractures occurs in 14,
folds in metasandstones and purallels the
weakly developed axial plane schistosily of
these folds (Fig. 57). The intersections of the
fractures {S.,) with the bedding detine wu ridge-
und-furrow lineutian which parallels the F,
fald axis and is referred to here as La (Fig.
59). These fractures have lirvely controlled
albitisition and the ¢mplacement of thin peg-
matites. They have also been planes of past-
alternation movement ulong which either thin
cuatings of fibrous hornblende or hrecciated
zones have developed.
Inspection of the metasediments in the
Middleton area shows that three schistosities
ure present. 4 well expressed axial plane whit
tosity in some mesoscale folds in fine grained
metasandstones can be seen in thin sections
to crenulate a poorly developed older schis-
losity that is approximately parallel {o bedding
(Fig. 60). Moreover, an examination of the
axtal plane schistosity surfaces. shows u con-
spicuous mica-streaking lineation tbat is not
parallel to the fold hinges (Fig. 613. The
presence of two schistosities in these folsis
= Mésosealé Fo totds are Wneommon. but gencrally oceur nenr jhe margins of pre-Fu pegmatite dykes
in jhe Petrel Cove Formation, No wllempl was made to measure their attitudes in such localities,
GEOLOGY OF TYPE SECTION, KANMANTOO GROUP (CAMBRIAN) 225
MN
+ Poles to So
8,= 200° ptunge 25 ® L)= cleavage/bedding
intersection
Vi Le=calc-silicate rod
MN elongation
MN
4 Poles io S,
o Ll
+ F, Fold axes 2
A FoFold axes
A,= 051" plunge 35° 8,= 118° plunge 50°
Poles to Sq
Pol
Poles to Sj * Hes: te Sg
dene
Ly
Lif
Fig, 6. Equal area projections of structural data. a-d, Tunkalilla Beach to Rosetta Head; e, Brown
Hill area; f, Middleton quarry.
22 RB. DAILY and A. R. MIT-NES
would normally identify the folds as secund-
generation structures. However, small scale Fy
folds in the Middleton quarry are characterised
by a poorly-developed axial-plane schistosity
(S) that overprints a
developed schistosity parallel to bedding. In
these folds. it is the older schistosity that con-
Wing @ prominent mica-steeaking limeation
whieh as similar in nature and orientation to
lig L,’ lineation recorded in metasediments in
ihe couslal part of the type section and in the
Brown Hill area In the Middleton quarry. Lhis
mica-streyking lineation is readily seen on the
surfaces of F, folds, where it is oriented at a
significant ungle to the intersection bewween
bedding and S, (Fig, 621. In view of these ab-
servations, we presently interpret the meso-
seale folds of uncertain uffinitics. os first-
pencration siructires that formed in metiasedi-
ments containing a pre-teetonic bedding pline
schistosity, and the accompanying prominent
micu-streaking lineation as by‘.
F,, olds were nol observed in the rather poor
exposures outside the Middleton quarry, o i
the small quatty expasures NW of Port Ri hat,
In these foculities, folds also hesitantly inter-
ptered as first-generation structures form the
dominynt sttuctueal clements. They are best
seen i the exposures to the north of the
Middleton quarry. They are inclined, and
plunwe jd steep angles towards the SE. The
uXial plane schistosity is a pronounced mica-
preferred urientalion which again overprints a
pourly preserved presumed pre-tectonic bed-
din-plune schistosity in some specimens, The
imerseation ot bedding with the axial-pline
sclisiusity is parallel to the axes of the folds,
However, a prominent micd-streaking lineation
oa schistusity surfaces pitches up to 20° from
the intersection between bedding, and the axtil-
plane schistosity-
S, Satiaces in phyllites of the Petre! Cove
Formation exposed inthe quarries NW of Port
Elliot and in the cuttings along the Crow's Nes!
road cxhibil a fine-scale crenulation. ‘The creou-
lalion appears to have a similar orientation to
the conspicuous mica-streaking lineation { pre-
sited L,') on $, surluces in metasandstones
in the Middleton urea, but ts interpreted a3 a
secomiu-poueration structure.
Theexposures of Middleton Sandstone along
the coasuine at Middicton are dominated by
F,, Structural clements. A proninent ridge-and-
lurrow linewtion (L.) indices a shallow
easterly-plunging fold. This ayrees with the
moderuicly well-:
orientalion of the fold axis of the only Fold
recorded along the beach.
The geometry of the structural elements for
the Middleton sree are given in Figs, 6f, Tae
und show the following: Poles to S, measured
in the Middleton quarry (Pig. 6F) do nut re-
flect the attitudes of firscgeneration folds, burt
define and F. fold axis estimated to plunge
towards LL&° at SO". This agrees. with the atti-
tudes of measured Fy fold axes and L,, ridye-
and-furrgw tinealions (Mig. 7a), In addition, it
is Consistent with the distribution of poles un
S. fractures in melasandstones.
The distribution of measured mica-streaking
hnewtions in the Middleton quarry indicates an
estimaled plunge towards 122° at 65°, whilst
the attitude of the related schistusity indicates a
stecply-plunging inclined style for folds herein
interpreted as first-generation structures (Fig.
7b).
The structoral elements Sy, and 1... measured
in the exposures at Midulleton Beach define an
F., told axis estimated to plunge at 28" tawards
NYO", and this is supported by the attitude of
one mesoscale F,, fold uxts (Fig. 7c).
In all other parts of the Middleton area,
presumed first-generation structures are domin-
ant, Poles to S, define an B, fold axis csu-
mated to plunge at 60° towards 142° (Tig,
7d). The rather diffuse distribution of S, poles
on the western side of the diagram may reflect
the influence of second-generation folds, ‘The
attitude of the F, fold axis is confirmed by the
distribution of mica-streaking lineations und the
attitudes of poles to the axiul-plane schistosity
(Piz. Je), and approximately coincides. with
the orientation of presumed first-generation
structures recorded in the Middleton quarry.
BL. STRUCTURAL RELATIONSHIPS OF THE
Encoun?tit Bar GRANITES
The contact between the Kanmantoo Group
metuscdiments (Petrel Cove Formation) and
the Encounter Bay Granites is exposed on
Rosetta Head and Wright |, bat is best
examined on Waiglit 1. because of the excellent
exposures there. Our observations of the rela-
tionships at this contact can be summarised is
tollows.
{) The marginal phase of the Encounter Bay
Granites is a coarse-grained meguerystic
granite, Megacrystte granite sheets of
variable thickness and grain. sixe form
apophyses from the main granite mass,
and ure laracly concordant with bedding
in the Petfel Cove Formation metasedi-
GEOLOGY OF TYPE SECTION, KANMANTOO GROUP (CAMBRIAN) 227
B= 18° Plunge 50°
Poles to So
o bg
4 F53Fold axes
¥ Limica streaking
4 Poles to $4
MN MN
o
B:=O90" plunge 28
B:=142° plunge 60°
» Poles ta So » Poles to Sg
on lg
A FoFoid axes
MN
B,= 142° plunge 60°
¥ Limica streaking
& Poles to S,
o Poles to Sg
Fig. 7. Equal area projections of structural data. a-b, Middleton quarry; ¢, Middleton Beach; d-e,
Middleton area excluding the Middleton quarry and Middleton Beach.
4
rm
x
ments TFig. 63). The granite sheets dre
murkedly boudinaged in places,
[h) Petrel Cove Formation metasediment, in
contaer with the megacrystic granite, hoth
on Wright I. and Rosetti Head, are lim-
inated metasiltstones, A paucity af porphy-
roblastic andalusite and cordierite schists
adjucent tn the contact compared with
their abundance at lower straliyraphie
levels Within the formation further from
the granite was noted,
(c) A prominent schistosity ($4) occurs in the
Petre] Cave Formation metusediments anil
@ pafallel schistosity occurs In the borders
and constricted portions of \he boudinnged
megaerysiic granite sheets (Fig. 64). The
same schistosity is variably developed
within the marginal metre of the main
granite mass. Within the affected granite,
the schistosity (Fig. 65) is defined by al-
ternuting laminae of recrystallised biotite
and quartz, which cnwrap latge potash
feldspar and pligioclase megacrysts. Al-
though recrystallised along their margins,
these megucrysts retain their original inter-
nal structures: some have been rotated.
The schisloxe gronile muy be regarded as a
protomylonite using the terminology of
Hiseins (1971). However, we believe that
the schistosity has been imposed during
regional deformation at a moderately high
temperature. The variable development of
the schistosity in the srinite along the
contact. 1ogether with its absence from the
internal parts of the pluton, gre duken ws
eVidence for the high yield strength of
the granite at this time, and suggest thit
the pluton possessed sigh bouy properties
during the regional deformation,
Metisciliimentury-rock xenoliths within the
megacrystic granite awoy [rom the con-
act ure devaid of deformation structures
and, with few exceptions. tectonically tm-
posed miga-preferred orientations. More-
aver. xenoliths containing andalusite or
corilierite porphyroblists have not becn
observed.
(el Structures younger than the S, schistosity
occur wilhin the metascdiments and
wranite sheets on Wright 1, and include
fine scale crenulations in the 5, schistosity.
Broad scale kink-folding of bedding add
thin eranite sheets has alsa been noted,
On the southern side of Wright 1., a cata-
clastic zone crosses the contact and has
deformed the S$, schistosity. Catactastic
(al)
Kk IALLY and A, Re MILNES
TOAtUIES are CONspicuoUS in the granite.
and are strikingly different from textures
in the schistose granite.
C, Summary
The observations anc meastirements of
structural elements indicate two malt phases ot
deformitiun of Kanmantoo Group metascdi-
ments in the type section. With the exception
of parts of the Middleton area, first-generation
structures ate dominant. However, second-gen-
eration structures are moderately well devel-
oped in the Petrel Cove Formation between
Rosetta Head and King Beach, and are domin-
ant in. Middleton Sandstone in the Muldleton
quarry and at Middleton Beach,
There ate several structural observations thut
will only be adequately explained after the
camipletion of a more comprehensive investi-
gation of the structural geology of the Kammun-
too Group than was attempted here, Such ob-
servations include the variable development of
structural elements in the type section, and the
variability in style aod urientation of first
generation folds. Notwithstanding, the data
presented herein provide an adequate basis for
an interpretation of the time of emplacement
of the Encounter Bay Granites in relation to
the structural deformations recorded in the con-
liguous Kanmantoo Group metasedimentary
rocks. The following observations are con-
sidered most pertinent to this discussion;
(a) The prominent schistosity that aceurs with-
in some parts of the megacrystic granite
along its contact with the Kunmuantoo
Group metasediments, and in boudinaged
grantle sheets: wilhin metasediments auja-
cent to the contact, is parallel 10 the S,
tchistoxsity in-the metasediments; and
metasedimentary-rock xenoliths that occur
within the meguerystic granite away from
the contaci. are. with few cxeeptions, de-
void of lectonically imposed mici-
preferred orientations,
Thus the Eneounter Bay Grnites ate con-
sidered to have been emplaced prier to the
main phase of first-generation deformation and
are, in (his sense, pre-tectonic, The texture of
the schistose granite indicules that the meyi-
crystic granite had completely crystallised prior
to the imposition of the $, schislosxity during
the main deformation phase,
Although second-generution structures have
overprinted the 8, schislosity in the metasedi-
ments and in ihe biotite-rich borders of the
granite sheets on Wright 7, their sporadic de-
velopment bas prevented ar) assessinent of their
fh)
GEOLOGY OF TYPE SECTION, KANMANTOO GROUP (CAMRBRIAN)
effect nn the granites an a broad scale. On the
other hand, second-geheration structures pro-
vide an important teference point in a discus-
sion of the relative ages of pegmutite and meta-
dolerite dykes. For example, hoth peematites
und metadolerites occur ag transgressive dykes
within the Petrel Cove Formation, and do not
exhibil first-generation structural elements,
However, they have heen folded and houdin-
aged during the second phase of deformation,
Ir fact, second-generation structures, inchiding
mesoscopic folds and an obvious crenulation
cleavage. are commonly hest developed immed-
dutely adjacent to these dykes, On the basis of
these observalions, the pegmatites and meta-
dolerites appear to post-+late the first-generation
folding and pre-date the second. Similar rock-
types idtrudiog melasediments further west
in the type section are probably of the same
age, although this cannot be confirmed because
of the iipparent absence of second-generation
siructures in these areas,
Metamorphism
Published studies on the metamorphic pet-
rology of the type Kanmantoo Group have
centred around the metasediments in the
vicinity of Rosetta Head. From that area
Browhe (1920) first recorded the presence of
indalusile und cordierite . Later, Bowes (1954)
subdivided these metuseciments into three
groups, namely quartz-blotite schists, anudulu-
site and cordierite schists, and albite and chlor-
ite schists. Both Browne and Bowes regarded
the metamorphic assemblages as a consequence
of the emplacement of the Encounter Bay
Granites. On the other hand, Offler & Fleming
(1968, p. 259) recognised “snowball” Internal
fabrics in (he andalusite and cordierite porphy-
roblasts in this area and suggested that “since
the Victor Harbor Granite does not appear to
have been emplaced forcefully. such fabrics
could only have been produced during a defor
tation phase before the intrusion of the gpran-
ites. As shown below cordierite is mainhy pre-
(o warly svn-F, whilst andalusite is late syh- to
post-F, and pre-F,,,
For that part of the type Kanmantoo Group
west of Tunkalilla Beach, Daily & Milnes
(T971u) concluded that the metamorphic arnde
lay between the almandine and slaurolite isa-
studs as defined by Winkler (1970+, and with-
in the andalusile-staurclite zone of OMer &
2349
Fleming (1968). Although our estimate: of the
prade still stuns, we realize that our inter-
pretation of the metamorphic history was over-
simplified. In the present context we have
uttempted to link the various mineryl assem-
blages to readily identifiable structural elements
in the rocks and thus deduce the retative time
relations between crystal growth and deforma-
tron in the fashion of Zwart (19631, Spry
(1963) and Offer & Fleming (1963), How-
ever, this method cannot give a measure of the
time between the various structural and zon-
comitunt metamorphic phases, nor can it allow
one to assess the time tiken for the essential
metamorphic reactions to produce the mineral
ussemblages seen today. Tt is most likely thr
the various recagnisahle metamorphic phases
discussed below are but part of a continuum,
It should be noted thal all the following ob-
servahions and the conclusions drawn from them
pertain only to rocks in the type section of the
Kanmantoo Group as specified by Sprigg &
Campana (1953) and jin our extension of the
type section into the region between Brown
Hill and Middleton Bench, which we regard
48 helonging to the uppermost part of the Kan-
mantoo Ciroup in the castern Mt. Lofty Ranges.
Within the Kanmantoo Group conspicuous
metamorphic mineral assemblages are uncom-
mon, and are confined mainly to thin cale-sili-
cate banifs and lenses and to satne pelitie inter
vals, All nictusediments except the phyllites re-
tain aspects of Iheir original detrital nature
despite the deformation and metymorphism,
The following assemblages of minerals? scem
to be representative of the vurinns rock types
wilhin this region:
(a) metasandstones and metasillstones—
quartz + plagioclase + biotite ++ musco-
vite = calcite + chlorite 7 garnet
scapolite = epidete;
(hd phythtes—
quariz — plagioclase + biotite + muses-
Vile = chlorite + varnct + andalusite +
cordierite;
(¢) carbonaceous and sulphide-rich phyilites—
quartz + plagioclase + muscovite +
pyrrhotite + graphite;
cale-silicates—
quariy | plagioclase + hornblende = gar-
net + chlorite + calcite = biotite + mus-
covite = pyrrhotite “= epidote.
(d)
* Offer & Fleming (196%) have reported fibrolite in many schists within the Mi Lofty Ronges. We
had nal seen thts mineral in our thin seetions until Dr. R, Ofer located am isolated patel in a thro
seciion of metasandstone collected 200 m east of Bollaparadda Beach,
134
AS uv fesult of a petrographic study of the
Kanmantoo Group rocks collected during our
traverses, ihe following scheme of progressive
mctamorphic crystallisation is envisaged:
A. Pre- te Syn-F, Metamorphic Crystallisation
Metamorphic elements attributed to this
phase of crystallisation inglude small quartz-
rich aygtexales, groundmass. biotite and mus-
covite, porphyroblasts of biotite, andalusite and
rare chlorite, and cordierite augen, the latter
commonly altered to a brownish-yellow clay
mineral determined hy electron probe analysis
as kuolinite.
BiotLe porphyroblasts occur with ground.
mass micas throughout the region, but the
quariz-rich aggregates seem to be restricted to
the remon hetween Tunkalitly Reach ard Bn-
counter Ray. Cordicrite is more severely restric-
ted, bein found only in some phvilite hands
in the upper part of the Balquhidder Forma-
tien, and mm the Petrel Cove Furmation neni’
Rosetta Head
Metamorphic clements of pre-F, age ore
uncommon but include many small quartz-tich
ugercgutes containing feldspar, opaques and
micas (Figs, 66 & 67), Their grain size is
always finer than that of the same minerals in
ike groundmass. They form augen which
generally contain a randem internal mica fab-
Tic and arc enwrapped by the well developed
mica schistosity. However, same quurtz-rich
uugen that have been observed with a poorly-
oriented mica fabric may be partly syntectonic.
Some cordicrite augen in the Petrel Cove Fors
mation exhibit a random inclusion fahrie und
represent pre-P, crystallisation.
Metamorphic elements formed during the
first. phase of deformation include the minerals
that define the S, schistesity, namely biotite
and. muscovite. Fer the most part, syn-F, bio-
tite and’ muscovite are fine grained, Medium-
grained biotite and chlorite parphyroblasts of
this age have been noted, 5, is also commonly
marked by u preferred orientation of opaque
mineral laths, for example Fig. 66. Many cor-
dierile wugen which are enwrapped hy $4 show
oriented inclusion fabrics, that are cither planar
and inclined at a significant angle to 8. or are
S shaped (Figs. 67-70). Such fabrics are con-
tinuous with the 8, schistosity in the enclosing
rock. Moreover, as inclusions within the por-
phyroblasts are finer than the same minerals in
the groundmass, we conclude that cordicrite
crystallised and was rotated during, the early
Stages of formation of 3, -
B. DAILY amd A. R. MILNES
Andalusite commonly occurs as S-shaped
poikiloblasts containing S-shaped inclusion-
trails that are continuous with the external S$,
schistosity (Fig. 71)_ Such poikiloblasts are the
result of late syn-S, crystallisation ond rotation,
In some cases the outer margins of poikilo-
blasts cut across this schistosily and must ihere-
fote represent post-S, crystallisation (Fig, 72)
H. Past-Fy Metamorphic Crystallivation
The major porphyroblastic crystallisation
appears to have occurred during the post-F,
and pre-B, static phase when andalusite, gur-
net, hornblende, biotite, chlorite, muscovite.
scupolite and epidote crystallised in rocks of
appropriate lithology. These minerals cut across
the groundmass S$, schistosity, but are de-
formed by the $. crenulation cleavage in rocks
in which this structure is developed, The fol-
lowing conspicuous minerals aid mineral
assemblages are characteristic of this meta-
morphic episode:
ta) calc-silicates—
hornblende “4 plagioclase +
chlorite + epidote,
metasanrstones and metasilistenes —
muscovite + garnet = chlorite + epidote;
(c) phyllites——
= jndalusite = chlorite =
garnet + scapolite + biotite.
In addition, {here are several miner com-
poncots such as tourmaline, some opaque
minerals and sphene which appear to have
crystallised across the groundmass $5 schis-
fosity, and are similarly attributed to this meta-
morphic episode.
Post-5, andalusite occurs both as over-
growths on svn-S, poikiloblasts (Fig. 72), and
as poikiloblasts that have overgrown the
groundmuss §, schistosity and have retained
relicts of this in the form of oriented opaype
laths and quartz grains (Figs, 73 & 74), In
addition, post-S, chlorite (Figs. 75 & 76) and
less commonly muscovite occur in phyllites and
metasiltstones throughout the type section as
Poikiloblasts that have grown across $4. Scapo-
lite occurs as small porphyroblastic clots that
appear to have grown across 5, within meta-
siltstones, 1 kr east of Coolawang Creck-
Porphyroblastic zoned garnets (Fig. 77) may
occur with minor muscovite, chlorite, epidote
and tourmaline in mcetasiltstones and metasand-
slones throughout the lype section, These have
eTown across the S$; schistosity. Such garnets
aré conxpicnous in the heavy mineral sand
suites along the southern coastline.
gatmec
(b)
muscovile =
GEOLOGY OF TYPE SECTION, KANMANTOO GROUP (CAMBRIAN)
Although an interpretation of the textural
relutionships im most rocks éiggest that the
crystallisation of cordierite preceded that of
undulusite, co-exisling post-F, cordierice
poikiloblasts and andalusite euhedra occur in
pelites within 1 m of a metadolerite dyke about
half way between King Beach and Rosetta
Hew. As structural evidence indicates that the
metadolerite intrusions are pust-F, but pre-F.,,
the cordierite-andalusite assemblage is inter-
preted as being the result of a local increase in
temperature along the dyke margins.
Post-F, culc-silicate segregations occur
throughout much of the region and possess a
weakly defined relict 8, schistosity that con-
trasts with the better defined schistosity of the
enclosing mietasedimeots. Hornblende poikilo-
blasts (erroncously identified optically as
actinolite by Duily & Milnes 1971a} occur in
the cale-silicates and, at best, exhibit only a
moderaic depree of preferred orientation
parallel co the relict S, schistosity. Some horn-
blendes. transverse to S, contain undeflected
remnants of that schistosity, especially opaque
faths. Zoned varnets occur in the cale-silicates
ws subhedral poikiloblasts that cut across S,
but these also have inherited the 5, schistosity-
Post-P, metamorphic minerals that occur in
peliles in the eastern part of the type section,
where the S., crenulslion cleavage is best de-
veloped, ure deformed by Sy. The cislocation
of chlorite and undalusite poikiloblasts along
zones commoniy defined by trains of opaguc
minerals grains (Fig. 78), are especially evi-
dent in pelites In the Petrel Cove Formation
in which Sy assumes the characteristics of a
stramn-slip cleavage,
Many pegmiutites within the type section are
composed of minerals that characterise the
post-F, mineral assemblages in the host rocks.
On structural evidence, such pegmatites are
post-F; and pre-Fy and represent the “sweat”
products concentrated in regions of low pres-
sure gtadient (Rivalenti & Sighinolfi 1971)-
Wall-rock alteration has been noted adjacent to
some of these pegmatites.
C. Conditions of Metamorphic Crysiallisaion
A petrographic examination of Kanmantoo
Group metasediments in the type section seems
to indicate a relatively simple sequence of
Metamorphic crystallisation, This sequence be-
gan with the formation of quartz-rich agere-
gates and the crystallisation of cordierite dunng
the pre- to easly syn-F, phase of deformation,
was followed by ihe late syn-P, crystallisation
of andalusite and chlorite and ended with the
231
post-F; crystallisatioi ot andalusite, chlante,
gamel, hornhlende, muscovite and scapulite,
Offler & Fleming (1968) regarded the meta
inorphic mineral assemblages Whtoughout much
of the Mount Lofty Ranges as churacteristic oF
low pressure, intermediate-type metamorphism,
With reference to the aluminosilicate data of
Newton (1966), they suggested that meta-
morphism in the areas of highest grade
occurred at pressures (Pigiai — Pang) between
3 and 4 kb and at temperatures in the vicinity
of 650°C, The metamorphic minerals in the
Kanmantoo Group metasediments in the type
seclion are certainly consistent. with conditions
of low presstire, intermediate-type metamar-
phism, but are representative of lower tem-
peratures und pressures than the assemblages
upen which Offler & Fleming based their esti-
mate.
The crystallisation of pre- to syn-F, cor-
derite and late syn- to post-F, andalusite in
metasediments in the type section suggests that
the conditions of metamorphisin at this time
Were consistent with the andalusite-cordierite-
muscovite subfacies of the amphibolite Facies
(Winkler 1965). Of several possible para-
geneses fnvolving the crystallisation of cor-
dicrite, the reaction—chlorite ++ andalosite +
quartz = cordierite + vapour (Sieferr &
Schreyer 1970) seems to be consistent with the
observed mineral assemblages, Based on the
experimental P-T conditions for this reaction,
and for the aluminosilicate stability as deter-
mined by Holdaway (1971), the crystallisition
of cordierite and andalusite in metasediments
in the type section probably occurred at pres-
sures below about 3 kb and al temperatures
estimyted to be less than 540°C (Milnes un-
pub. Ph.D. thesis) -
Discussions and Conchusious
As a result of our studies on the upper part
of the type Kanmantoo Group, several facts
emerge from which w nurnher of conclnsions
can be drawn additional to those given in Dally
& Milnes (197tay-.
A. Srratigraphile Relationships
In no instance throughout the whole of the
coastal sections have we been unable to ascer-
lain bedding. Transposition as described by
Talbot & Hobbs (1968) is confined to the
vicinity of Petrel Cove. Most outcrops retain
their gross sedimentary features despite the Lec-
tunis and metamorphism, This is readily dis-
cernible from the accompanying fignres,
44
232
Vhere appears to be conformity between all
the fermations muking up the Kanmantoo
Group as sec out in Table 1. Phe validity ot thy
straligtaphic scheme is enhanced by the oceur-
reace of a virtually identical sequence on Dui-
ley Peninsul, Kanguroo 1, (Daily & Milnes.
I971b, 1972b}. As Our investigations on. this
ishand ore as vet jacomplete there is still room
for an upward continuation of the sequence
beyond the Middleton Sandstone. The latter
constitutes the youngest purl of the Kanmantoo
Group in the Mt. Lofty Ranges, Lt is worth
nog that in the original definition of the
Kanmantoo Grolp, Sprigg & Campana (1953,
p. 14) stated that “the upper boundary of the
Kanmantoo Group remuins undefined”.
B. The Kungarvoian Movements and Karrnan-
too Group Sedimentation
All the original sediments making up whe
Brown Hill and Wattaberr) Sub-groups in ihe
lype section were clastics which ranged in grin
size From clay to pebbles ut least S em in dia-
meter limestones were absent. Indeed. car
bonate shows up only as pebbles in conglomer-
ates or pehbly sandstones. However, its former
presence as iin accessory is probubly indicated
by epidote-rich bunds in the Midelleton Sand-
sions innit ws the thin calc-silicate sesrecations
and hands in all the other stratigraphic units
within hath sub-groups,
Metasandstone is by far the dominant
lithology throughout most of the Kanmuntoo
Group. The sands were essentially immature
all contained vurying amounts of clay and silt
Lmud) as matnx, Quartz and subordinate feld-
spar were the dominant sand-sized particles ind
were derived mainly from the older Precam-
brian Urystulline basement,
The rapidity of deposition may he giuged
mo only ty the immaturity of the sunds but
alsy trom the dominance of pirallel lamination
within the metasandstones. We Interpret this
type of bedding as a consequence of the high
flow regime that prevailed through much of the
depositional history of the group, Intervals with
cross-bedding and especially current-ripple
lamination are common sind represent deposi-
tion at lower flow power as has been estub-
lished by Simons ef al, (1965), Guy er al,
(1966) and Williams (1967). Aflen (1970,
Figs, + & 6), using some of these experimental
data, hus shown the possible alternative
sequences of sedimentary structures that may
form for yarying graity size with decreasing
flaw power at the time of depasiunn,
‘The sands were taid dowat by currents, which
&. DAILY und A. R. MILNES
over long periods of Lime were largely unidirec-
tional, Measured current directions from sedi-
ments infilling low amplitude scour-channels
wihin the Brown Hill Sub-group indigule dis-
(ribuiion of sediment by currents flowing
generally from the NW quadrant. This is simi-
far lo current directions established for the
uoderlying Inman Hill Sub-group, We have
been unable to detect a decrease in pebble size
towsrds the east for conglomerutes in the
Brown Hill Sub-group, though such aw varalion
is discernible in the underlying Inman Jill Sub-
group. For instance, conglomerates in an aver-
turned sequence of metasandstones at Penne-
shaw, Kangaroo T., contain clasts up to 30 cm
in diameter. In correlative conglomerates in the
uppermost parts of the Tapanappa Formution
near “Tunkalilla Beach, the maximum pebble
size is 7 cm. Uhe grealer distance of transport
for these pebhles is jlso reflected in the lower
proportion of carbonate clasts relative to other
clast types,
The persistence of lenticular conglomerates
und pebble bunds into the upper levels of the
Balquhidder Formation. indicates that the neat-
hy tectonic fands, elevated in response to the
Kingarooian Moyements, periodically contri.
buted gravels to the basin (fur discussion on
these movements sec Daily & Milnes 197 1a, p,
20). Evidence, mamly from Kangaroo L and
Yorke Peninsula, indicates that the pebble
suites were derived from the erosion of a strati-
graphic <equence involving crystalline base-
ment and its uncenformable cover of Lower
Cumbrian sediments, mainly carbonales.. Within
the Balquhiikler Formation the buses of the
sands are sharp. No Atte casts, tool murky or
groove casls were located. The tops of sands
tend tn be similarly abrupt.
In contrast to the sandstones of the Brown
Hill Sub-group. the sundstones constituting the
Middleton Sandstone are much cleaner ane in
Utis regard ure similur to those of the Back-
stairs Massage Formation. They are very well
laminated and the »bundant medium-scale
cross-bedded sets indicate current directions
almost invariably from the west, although some
reversed directions are known. Slumping is
common and always down the fore-set direc-
tron. We do net interpret the slumping as a
consequence of the slope of the basinal floor at
the time of deposition, More likely, it has
resulted from the mass movement of the highly
unstable woter-laden (thixotropic) fore-ets,
perhaps due to current drag, The Middleton
Sandstone on Dudley Peninsula, Kangaroo ,,
GEOLOGY OF TYPE SECTION, KANMANTOO GROUP (GCAMBRIAN}
also conustenily shows fore-sets and slumps
thrected cowards the eust,
Load custs on the bases of fpetasandstones
ure common, Occasionally, where soft sediment
formation was more intense. pseudo-nodules
(Macyr & Antun 1950) were formed aid in
extreme cases isolation from the overlying
parent sand was effected, thus permitting the
balled-up sund to sink into the underlying
muds, Exampics of all these structures can be
secn in the Rosella Head area and were figured
by ‘Talbot & Hobbs (1969),
The Kanmantoo Group pelites above the
Tapanappa Formation frequently show aod
sedimentary structures, mainly iamination or
rippled features. The latter range trom starved
ripples, indicating insufficient silt tu fine sand
in the transported fraction ta form a continuous
bed. to sets of climbing ripples. “where sedi-
ment deposition occurs simultaneously with
continuing irinsport in suspension and as bed-
Joad” {Allen 1970, p. f) in the lower flow
regime. The latter structures are indicative of
very high rates Of deposition.
Convolute lamination is not common bul
where it occurs it can efter be traced along
strike for the full length of oulcrup, thus indi-
cating the widespread naturc of the events
which led to its formation,
According to Sprigg & Campana (1953,
Pp. 12) the dominant facies of the Kanmyntoo
Group “are comparable with the Alpine
Flysch"”. Dally (1956, p. 139) stared “It is
believed that shoreline conglomerates, sana-
stones and shales as see in the Kangaroo
Island Group grade seawards into the ‘Aysch-
like Kanmantag sediments which were
deposited in areas of rapid subsidence’. There
is mo doubt that the characteristic alternation
of Metasandstones and pelites as seen in the
Rulquhidder Formation is: ‘“flysch-like’, How-
ever, in no respects can we regard this as o
turbigite facies. For instance, nowhere have we
seen the five-fold division of internal sedimen-
lary slructures which characterise the ‘cam-
plete” turbisile of Bouma (1962). Ta fact, pro-
longed search has fuiled to find the truly graded
and structureless bisal interval which features
so prominently in Bouma'’s “turbidite facies.
model". The tyct that somie grading may occur
within the well-luminwted and ripple-bedded in-
tervals is no criterion of deposition by Lur-
bidity currents because comparable grading
oceurs in sequences deposited in a number of
enviranments, marine and non-marine. In his
critical review of the turbidite problem Vin
235
der Lingen (1969, p. 10) also used the term
“Mysch-like” to cover “alternating, sequences
reuching thicknesses of geosvneclinal magnituule,
bul fackiny typical “turbidite characteristics’ ”.
In our Opinion it is wrong to cquate “fiysch-
like™ sequences with turbillites, be they either
proximal or distal (Wiulker 1967, Table IL, p.
32), or Nuxoturbidites, (Dzulinski er af, 1959).
What must be proved of course is that the
sequences concerned are the products of tur-
bidity currents before they cah be called dur-
bidites. In addition, the terms flyweh and
furbidites conjure up in many people's iminds
the notion of deep water environments, Already
there are many instances Where such iclews are
incompatible with the evidence. for cxample
the discovery of the footprints of birds in some
meas of flysch (Mangin 1962).
The interpretation af the depositional envir-
onment of a sedimentury sequence may be de-
duced only after critical cxuminalion of all the
available cvidence, Rven then no finality need
be reached, In the present context we visualise
the following:
(a) Because of the great thickness of Kanman-
too Gtoup metasediments. subsidence
must have been extremely rupid, Basin
development was probably controlled by:
faulting to account for this.
‘The nature of must of the racks shows thal
sedimentation was rapid. Possibly ueposi-
Lion kept pace with subsidence. There is
no evidence of lemporary emergence, such
as mud-cracks. However, these would he
hard to-Jocate in this metamorphic terrain.
(c) Within the Balquhidder Formation, low.
amplitude scour-channels are common in
the thicker sand intervals. The associated
low angle cross-hedued sets (generally less
than | m thick) are typical of deposition
in an aqueous environment. A marine en-
vironment is suggested by the prevalence
of the higturbated tops of some of the
sandicr bands. Severe restriction giving
rise to unfavourable environmental condl-
tions could be invoked to explain the pre-
sumed absence of a benthoni¢ shelly fauna
in the sands. However, such 4 restriction
may be more apparent than real, for in
view of the rapid rate of deposition, the
basinal waters would have contained abim-
dant fines in suspension which may have
heen deleterious to the presence of a Cam-
brian shelly fauna. Assuming a marine en-
vironment, the lack of. reworking of the
sediments indicates that either subsidence
fb)
34
wus rapid enough to prevent reworking Ur
that sedimentation look place at a depth
below effective wave base. This could have
heen quite shallow (McCave 1971}.
The high incidence of medium scale cross-
bedding and the greater maturity of the
sands of the Middleton Sandstone scem
lo indicate that it was deposited in a
shallower environment than the Balqu-
hidder Formation, It is almost certainly a
shallow water deposit.
In the absence of autochthonous foxsil
assemblages it is impossible to give any objec.
live depth indicators purely on the basis of the
known sedimentary structures. All we can sug
gest here is that the Middfeton Sandstone is a
shallow water deposit, which because of rapid
subsidence escaped reworking, Rocks of the
Balquhidder Formation were — probably
deposited in somewhat deeper water but there
is mo yuarantee of this, We certainly do not
envisage ji deep water origin for it, because un-
metamorphosed sediments of similar facies and
ef shallow Water origin occur on the northern
coast of Kangaroo J, Possibly the deepest water
is indicated by the finely laminated blue-black
carbonaceous and sulphide-rich phyllites indi-
caling stagnant bottom conditions, However,
tven here avery shallow environinent of de-
position Is possible. All that is needed to pro-
dulce sediments of this type is a Call off in the
deposilion of coarser ¢clastics, a restriction of
circulation, and a supply of abundant organic
niatter to produce the carbon and sulphides
under strongly anoxidising conditions.
a
C. Relativuship of Metamorphism und Strtec-
jure to the Emplecemeth of the Encounter
Bay Granites
Our observations suggest that the Encourter
Bay Granites in the Encounter Bay ares were
emplaced prior to the main phase of first
generation deformation, Moreover, metamor-
phism in this area began with the pre- to eany
syn-F, crystallisation of cordierite and quartz-
rich pgeregates in metasediments of approp-
rule composition, continued with the late syn-
F, etystallisation of andalusite and chlorite,
and ended in the post-P, structurally static
phase during which the crystallisation of an-
dalusite, chlorite, garnet, muscovite, hornblende
und scapolite occurred in metasediments of
uppropriate composition,
Cordierite is restricted to some pelites in
the upper part of the Balquhidder Formation
and in the Pewel Cove Formation along the
Coustal section between Encuunier Bay and
B, DAILY und A. R, MILNES
Pursons Beach and its crystallisation records
the highest grade of metamorphism in the
revion, It occurs as altered augen in phyllite
bunds that crop out in the headland at the
easiemn eng of Parsons Beuch; it has not been
recoghised in racks to the west of this locality.
nor in rocks in the Brown Hill to Middleton
arex, Andalusite o¢curs mainly in some pelites
in the upper part of the Balguhidder Formation
and in the Petrel Cave Formation slong the
coastline east of Newland Head in association
with cordierite, However, ture altered post-F,
poikiloblasts that occur further west along the
cogstline, for example in Tapanappa Forma-
tion metasediments near the western end of
Tunkalilla Beach and at Tunk Head. are con-
sidered to have been composed of andalusite,
Their occurrence 1 consistent with the presence
ot andalusite poikiloblasts in Tapanappa For-
mation metasediments that crop out along [the
northern coast of Dudley Peninsula, just cast
of Penneshaw (Daily & Milnes, unpublished
observations). Minerals sich ag homblende
und garnet, which are characteristic of cale-
silicate lithologies, oceur throughout the type
section, Scapolite however, is confined to cer-
tain pelites that occur 1 km east of Coolawang
Creck.
We have mapped a magnificent development
of syn- to post-F, metamorphic minerals such
as andalusite. hornblende, scapolite and garnet
in Adelaide Supergroup metascdiments which
occur within a thrusted fault block along the
north coast.of Dudley Peninsula, approaintuicly
12 km from the Encounter Bay Granites at
Cape Willoughby (Daily & Milnes 1971b,
1972b). However, there is a conspicuous
paucjly of these minerals itt Kanmantoo Group
metasediments closer to the granites. Thus,
Kanmantoo Group mefasediments in this area
appear, on the whole, to he ef such composi-
tion (for example deficient in alumina and
lime) that they fail to contain metamorphic
assemblages thal are indicators of metamorphic
grade.
The entire type section of the Kanmantoo
Group, as well as the section on Dudley Pen-
insula, occurs within the regional andalusite-
staurolite zune of meiamerphism according to
Offer & Fleming (1968). The presence of
plagioclase of oliguclase-andesine composition
in the metascdiments we have examined is con-
sistent with this grade uf metamorphism. Thus.
the paucity of andalusite in all but parts of the
Balquhidder Formation and Petrel Cove For-
mation west of Petrel Cove is best interpreted
GEOLOGY OF TYPF SECTION, KANMANTOQ GROUP (© AMBRLAN)
as die © a scarcity of metasediinents of
uppropriate hulk composition. The restricted
occurrence also of cordierite mav be due Lo the
limited presence of metasediments of auitable
composition. However, the canspicuuus devel-
opment of large post-F, cordierite porphyro-
biasis near a post-F, metadolerite dyke intrid-
ing metasediments. on the const about half-way
between Petrel Cove and King Point suggests
that varied proximity to a heat source. such as
the Enoounter Bay Granites, may also have in-
fluenced ita development. The absence of cor-
dierite and andalusite and other porphyrahlas-
tic minerals in Balquhidder Formation and
Petre? Cove Formation metasediments in the
Brown Hill to Middleton area. compared with
their abundance in metasediments In the same
stratigraphic position along the coastline west
of Encounter Bay, is patticulariy difficult to
nceount for. It may be the resule either of
inappropriate bulk compositions af metasedi-
ments in the Brown Hill to Middleton area,
pethaps due to facies changes along strike From
the coastal section, or the effect of conditions
af significantly lower metamorphic grade. In
view of the occurrence of plagioclase of olizo-
clase-andesine composition in the melascdt-
ments in the Brown Hill to Middleton area, the
former sugvestion is favoured,
Thus. the relationships hetween structural de-
formation, metamorphism and the emplace-
ment of the Encounter Ray Granites in the En-
counter Bay area, as deduced from field aid
pelrographic evidence. can be envisaged as
Follows:
(a) Regional folding, was initiated, probably
with the development of broad-scale Folds
and the development of an incipient S,
schistosity,
(b) The Encounter Bay Granites were em-
placed, and simultancously incorporated
abundant metascdiment xenoliths, the
majority of which do not display an im-
posed mica-schistosity, Cordierite and
quartz-rich augen crystallised in metasedi-
ments. of appropriate composition adjacent
to the granites, in response to locally ele-
vated temperatures,
(c) The main phase of F, folding, a regional
event, deformed the marginal granite
phases which had already completely
erystallised, and brought to completion the
development of the penetrative $, fabric
in the Kanmantoo Group metasediments.
Andalusite oad chlorite porphyroblasts
345
erystallised in these metascdiments on a
regional scale as a result of suitable P-T
conditions.
Saniftcant porphyroblaslic yrowth of
metamorphic minerals. occurred during
the post-F; static phase, Post-S, snddalu-
site crystallised us Overgrowths on syn-S;
porphyroblasts, amd us poikiloblasts ad-
jacent to the post-S, striped layering (S.).
Pegmative and ametudolerite adykes were
emplaced.
fe) A second phuse of folding (F.,) deformed
pre-existing structures, including 5S, and
the post-S, “striped” layering, and was
responsible for the development of a cren-
ulation cleavage and a strain-slip cleavage
S.. The main development pf Fu struc-
tures occurs in the eastern part of the type
section, Evidence for metamorphic
crystallisation during or post-dating the
second phase of deformation has not been
observed. However, albitisation and the
crystallisation of fibrous hornblende has
occurred along the §, fracture cleavage,
This 15 believed to be associated with the
widespread fate stage glbitisation (Dasch
er al. 1971) affecting the Encounter Bay
Granites. As well, we have observed tour-
maline filled fractures with orientation
similat to thal of S.,
td)
Acknowledgements
The expenses relating to this work were de-
frayed hy a Grant made to the University of
Adelaide by Beach Petroleum N11 We wish
to thank Mr. R. C, Sprigg for his continuing in-
terest in the project and for arranging the
Beach Petroleum Grant.
We wish to thank Dr. R. 1, Oliver for
his criticigm of the manuseript und for his ¢um-
ments on the metamorphic textures; alsa Dr.
M. A. Etheridge, Dr. A. W. Kiceman, Dr.
P. DB, Fleming, Dr, KR. Offler and Mr. R. G.
Wiltshire for discussions in the field and cam-
ments on an early draft of the structural and
metamorphic sections of the paper.
Tn addition we wish to thank the Drafting
Section, C.S.1.R.0. Division of Soils. for the
preparation of the coloured maps und Profes-
sor J. L. Talbot for the use of his large thin
sections of Petrel Cove Formution metssedi-
ments.
736
B. DAILY and A. R. MILNES
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Fig.
Fig.
Fig,
Fig. <
B. DAILY and A. R. MILNES
Well laminated metasiltstoncs within the Tunkalilla Formation, south sidé of Tunkalillu
Creek, Scale in mm (total length 7.5 cm).
. Worm casts etched ott by sand blasting of metasandstones, Bollaparudda Beach.
Well laminated metasandstones occupying a cut-and-fill channel in Balquhidder Kornvation,.
eastern side of Tunk Head. Hammer Iength 28 cm,
Tectonically deformed load casts and associvted flame structures, on the base of a meta-
sandstone, about 0.2 km east of Tunk Head.
Climbing ripple-trains developed in metasandstones jiterbedded with mtetasiltstones, Same
locality us for Fig. 12.
An erosional channel cut into well laminated metasandstones, sume locality as for Figs,
9 .& 10. Channel was filled with shale chips and sands. The hammer is lying against the
bioturbated metasandstones shown in Figs, 9 & 10.
Cale-silicate rods developed in metasandstones and plunging parallel to the fold axis; same
locality as for Figs. 12 & 13.
Thin planar- and irregular-shaped segregations of calc-silicates developed roughly parallel
to the hedding in meétasundstones; same locality as for Fig. 15.
Well bedded and cleaved metasandstanes in the trough of the Tunk Head syncline about
0.6 km west of Tunk Head. Note the well developed sigmoidal cleavage (outlined), ‘The
eavernotis weathering of the rocks is a common feature along the coast of Fleurieu Penin-
sula and on Kangaroo I.
Thick hands of well-banded metasandstones interbedded with thin laminated phyllites,
trough of ‘Funk Head syncline. Note the cross-cutting pegmatiti¢ veins fo the tight of the
figure, 1.8 m tall.
View of top surface of a coarse-grained metasandstooe bed showing smeared out sedimen-
tary structures, Lineation (hammer handle parallels it) is due to a cleavage/bedding inter-
section; same bedding surface as that shown in the bottom left corner of Fig. 18. Sedimen-
lary structures within the central party of the bed are relativcly undistorted by the
cleavage. Only in the uppermost few centimetres of the bed is the cleavage sharply refracted
thus producing marked distortion or iecionic smearing of the sedimentary structures.
Beds: of metasandstones alternating with thinner intervals of less resistant phyllites and
metusilistones; centre of Tunk Head syncline.
Tectonicully deformed cutrent-hedded (rippled) and laminuted metasiltstones interbedded
with phyllites about 30 m west of Tunk Head. Note the abundance of thin ptygmatic pee-
matilic veins,
Sulphide segregstions ane veinlets within a 2 im thick blue-black phyllite/metasundstone
interval, narrow pitch immediately west of Tunk Head.
Fallen block of Balquhidder Formation metasandstone displaying # thin bund of small-
ae conglomerate about 0:4 km east of Bollaparudda Beach. Hammer head lying in plane
of bedding.
Well-bedded coarse-grained metasandstones within the Balquhidder Formation. Note small
granules and pebbles recurring up through the sequence. Lecality abont 0.5 km east of
Bollaparudda Beach. Width of hammer head 17 cm.
Bright orange-red coloured zones of feldspathization developed along closely-spaced frac-
tures within metasandstone beds, 1.1 km eesti of Rullaparuddy Beach. Note paucity of
tensional fractures within the phyllite interbed compared with their greater abundance in
the overlying and underlying metasandstones, However, feldspathised zones do occur along
some of the frachires. cutting the phyllites. Feldspathized zones also occur to the west of
Mollaparudda Beach.
Fig. 3
Fig. 2
Fig.
Fig.
Fig. 3
Fiz.
Fig-
Fig
Fig.
Pig.
Pig,
Fig.
Fig.
Fig.
34.
Lint
La
3h.
4.
38.
2. 41,
GEOLOGY OF TYPE SECTION, KANMANTOO GROUP (CAMBRIAN| 239
Pegmatites infilling tensional fractures in metasandstones of the Balquhidder Formation 0.7
km east of Coolawang Creek. Note the warping and feathering of the fracture infills.
WellJuminated metasiltstones overlain by current-bedded sets of ripple trains 1.2 km east of
Coolawang Creek, Hummer handle 15 cm Jong,
A_ prominent rib-and-lurrow lineation on the sole of a thick metasandstone bed 0.7 km west
of Parsons Beach. Boat for scale.
Conyolute lamination in metasiltstones west side of Parsons Beach. Coin 2.3 em in dta-
meter,
Small channel cut into a pebbly metasandstone and filled with cross-bedded and laminated
metasandstones, Adjacent to beach on western side of Newland Head.
Fluggy outerops of metasandstones with thin phyllite partings on the western side of New-
land Head. A more massive band of meélasandstone is visible on top of the exposed
sequence.
View looking SW along strike towards Newland Head. Sequence cansists of steeply-dip-
ping metasandstones and interbedded thin phyllites. Included also is a blue-black curbonu-
ceous and sulphide-rich phyllite,
View looking towards Rosetta Head showing the high cliffline cut in the Balquhidder For-
mation, Note the lower cliffline cut in Petrel Cove Formution between King Point (extreme
right) and Rosetta Head.
Pen (14 cm long) is parallel to bedding im metasiltstone bands interbedded in knotted
potphyroblastic schists just below top of Balquhidder Formation, about 0.6 km west of
King Point. Note the pale coloured strain shadows around the aagen aligned in the cleavage
which dips away to the left.
Relationships of various layerings in porphyroblastic andalusite schists occurring near the
base of the Petrel Cove Formation, (Sq} is bedding, (S;) is an augen layering in the
cleavage, (S,) [referred to as (3.) in text) is a pale coloured “striped” layering. Note also
the “pinch-and-swell” quartz-feldspursmicu pegmatilic layers developed parallel ta the
“striped” layering and along which dislocation has occurred. Pen for scale.
A small synclinal fold in andalusite schists plunging towards 210° at 25°. The bedding,
well expressed by the folded darker layers, is also parily expressed by some andalusite-rich
layers, Some steeply dipping fine "striped" layers can be seen cutting the bedding in the left
of the photo. Tip of pen for scale.
Extremely well-laminated metasiltstones in the lower part of the Petrel Cove Formation
about 0.2 km west of King Point.
Current-bedded ripples and other sedimentary structures in well-bedded metasilistones,
Petre! Cove Formation, King Point.
Deformed load casts, their attendant flame strictures, and ripple-trains in metasandstanes
in the Petrel Cove Formation, 0.75 km SW of Rosetta Head wharf. Note thin “striped”
layers: culling the deformed sedimentary structures. Two pale coloured ribs of coarser sand
are elongated towards 210° in the direction of the Fy fold axis, The “striped” laycring is
post Fy as at cuts deformed sedimentary structures. Pen is 14 cm long.
Deformed load. casis 10 m SW of Fig. 39. Kedding dips shallowly to right. Note that the
bedding has been slightly offset by shearing parallel to the cleavage. Noie also the broad
stripe, tight of the coin (diameter 2.3 cm), dips ala slightly lower angle than the cleavage.
The stripe has a thin central dark biotite layer margined by broad pale coloured stripes in
itrn margined by selvages of biotite.
View looking SW at outcrop of shallowly-dipping, well-bedded, fine-grained metasand-
stone interbedded with softer and darker coloured andalusite schists, 0.75 km SW of Rosetty
Head wharf. All beds are cut by a white “striped” Jayering.
240
Fig.
Fig,
Fig.
Fig,
hig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig,
ig.
Fig.
44,
45.
ig. 46.
47,
48.
49.
54.
56.
57,
58.
B. DAILY and A. R. MILNES
Note refraction of discontinuous white “striped” layering in schists. steeper in metasand-
stones. Same locality as Hig, 41.
Refracted “striped” Jayering (partly discontinuous) sloping to left and crenulation cleavage
(almost yertical) in andalusite schist, Same outcrop as for Figs. 41 & 42. Nate that the
“striped” layers are slightly folded in some of the andalusite schist layers. The crenulauon
cloavuge (Ss) is axial plane to these folds. No obvious folds occur in the overlying fine-
grained metasandstone which dips to left at a shallower angle than the “striped” layering,
Same outcrop as in Fig. 41. Note the refraction of the “striped” Isyering as it crosses the
andalusite-deficient bands. There they are thickened,
Feathering of “striped” layers in metasandstones of Petrel Cove Formation just west of Petre!
Cove.
“Striped Juyering markedly refracted in passing from metasandstones into interbedded
andalusite schists, Bedding dips steeply to right and occurs on the limb of a fold plunging
towards 205° at 25°. Note that there is a crenulation cleavage axial-plane {0 small-scale
Fs folds confined to the “striped” layering within the andalusite schists, Locality on western
side of Petrel Cove.
Andalusites preferentially developed along some of the “striped” Jayers west of Petre! Cove..
Lens cap 7 cm in diameter.
View from 1.3 km SW of Brown Hill Jooking towards Rosetta [Tead. Note metasandstones
of Balquhidder Formation in foreground sloping down towards lower ground on the ex-
treme left which is underlain by phyllites of the Petrel Cove Formation, ‘This is com-
Parable with the geomorphology to the tight of Rosetta Head just below horizon.
Cross-bedded and welltaminated metasandstones of the Middleton Sandstone near western
extremity of outcrop, Middleton Beach. Scale 11 cm long.
Pale coloured epidote-rich bands in well-bedded fine-grained Middleton Sandstone near wes-
térn extremity of outcrop, Middjeton Beach.
Epidote-rich segregations or nodules occurring parallel to bedding in the fine-grained
Middleton Sandstone, near western extrentity of oulcrop, Middleton Beach.
Pale coloured epidote-rich layers outlining uw markedly croxs-bedded melasandstone unit
(300 om thick) i the Middleton Sandstanc, Middleton qnarry, Sequence is part of the
steeper limb of the Fe fold illustrated in Fig. 57.
View of S, surface in Petrel Cove Formation andalusite-cordierite schist. showing the inier-
section between S; and bedding {I4) and a lineation (T.) defined by the elongation of
andalusite and cordiernte auyen. Length of (Ly) arrow 14 cm.
Fy fold in bedding and in a thin metamorphic pegmatite, Petrel Cove Formation, along
coast between Petrel Cove and King Beach. Note refolded F; folds. Coin 2.1 cm in dia-
meter.
Small-scale Fy folds in bedding in Petrel Cove Formation metasiltstane refolded by Fo fold.
Schistosily, oriented approximately EW in photograph, is Ss. Same locality as Fig, 54. Coin
2.1 em im diameter,
“Striped” layering in laminated metasiltstone in Petre! Cove Fortnation metasilistone, same
locality as in Fig. 41. Note median biotite- and/or quartz-filled fractures, and biotite-rich
margins of stripes. Coin 2.3 cm in diameter.
Lurgé-scale. SE plunging Fo fold in Middleton Sandstone, Middleton quarry. Light coloured
layers parallel to bedding are cpidote-tich, Promincnl fracture cleavage axial plane to fold
has controlled albitisation (light coloured cross-cutting zones). Figure (arrowed) 1.3 m tall.
Tight small-scale F, fold in Jaminated metasandstones on limb of large-scale Fa fold in
Middleton quarry. Scale on right 7,5 ¢m. long.
Fig.
Fig,
Fig.
Fig
Fig.
Fig.
Fiz.
Pig.
Fig
Fig.
hig.
60.
al.
AK
f6,
47.
a8,
69,
TI.
GEOLOGY OF TYPE SECTION, KANMANTOO GROUP (CAMBRIAN) 24|
Ridge-and-furrow lineation (Lo) defined by the intersection of bedding and fracture cleav-
ge axial plane to Fy folds, Middleton quarry, Scale provided by 5 cm wide compass.
A large thin-section of mesoscale folds: in which w well-developed flaring axial-plane schis-
tosity (S-fold) overprints a pooily developed schisiosity (S) parallel to bedding; Middleton
quarry, Scale cm graph puper.
A specimen of mesoscale folds in bedding from which the section in Fig, 60 was cul, Nole
fald hinges (1-fold). and conspicuous mica-stréaking lineation (I.) developed on. schistosity
axial plane to folds (S-fold); Middleton quarry. Bar scale represents 4 cm.
Small-scale F., fold showing ridge-and-furrow fineation (Le), which is defined by the inter-
section between bedding and the fractufe cleavage axial plane to the fold, Note the mica-
streaking lineation (I.) that occurs on the moderately well-developed $) schistosity
approsimately parallel to bedding; Middleton quatry. Bar scale represents J cm.
Thin concordant granite sheet intruding, laminated metasiltstones of the Petrel Cove For-
mation, Wright fT. Pen 14 ¢m long.
Boudinaged concordant granite shect in Petrel Cove Formation metasiltstones, Wright I.
Note well-developed schistosity in margins and constricted portions of sheet. While spots on
rock surfaces are small marine gastropods, Hammer length 28 cm.
Surface of slab of schistose megucrystic granite. Schistosity, defined by alternating biotite
and guariz-rich laminue. enwraps feldspar megacrysts (light cotoured megacrysts ate plagio-
clase, darker coloured megacrysts are pofash feldspar). Quuriz megacrysts did not survive
the deformation. Specimen collected from main granite mass on NE side of Wright J. within
I m of contact with Petrel Coye Formation metasilistone. Scale in mm.
Pre-S; quartz-rich augen enwrapped by Sy schistosity in thin section of Balquhidder For-
mation porphyroblustic schist SC97", 200 m beyond eastern end of Waitpinga Beach. Note
the preferred orientation of abundant opaque laths in .S;. Transmitted plane polarised light.
Bar scale represents 0.10 mm.
Pre-S; cordierite augen, now altered to kaolinite, enwrapped by gromndmass §) schistosity.
Thin section of Petrel Cove Formation porphyroblastic schist TI30A. Transmitted plane
polarised light. Bar scale represents (0.50 mm.
Field in Fig. 67. Crossed polars. Note fine-grained size of quartz inclusions within altered
cordicrite augen compared with grain size of quartz in groundmass. Bar scale represents
0,50 min.
Syn-S; cordierite poikiloblasts, now altered to kaolinite. Thin section of Petrel Cove Forma-
tion cordicyite schist SC106A. The poikiloblasts exhibit an oriented internal fabric which is
continuous with the external S; mica schistosity, and have been rotated relative to it. Note
post-S; muscovite Juth (M). Transmitted plane polarised light. Bar scale represents 0.50
myitl.
Field in Fig. 69. Crossed polars. Note fine grain size of quartz inclusions in altered cor-
dierite augen compared with grain size of quartz in groundmass, This suggests that cor-
dierite crystallised during early syn-S; phase. Absence of curved internal fabric suggests
cordierite porphyroblasts. were rotated bodily with respect to external 8; schistosity after
crystallisation was completed. Note post-8,; muscovite lath (M). Bar scale represents 0.50
mm,
Syn-S;, S-shaped andalusite poikiloblast (A), with S-shaped internal fabric. Internal fabric
continuons with external groundmass. $1 schislosity. Note post-Si overgrowth on beltam
right of poikiloblast just lefp of post-S; chlorite lath (c). Thin section of Petrel Cove For-
ebnath andalusiie-cor dinrite schist T108. Fratismitied plane polurised light. Bar scale repre-
sents 0.50 mm.
= Numbers prefixed by SC refer to rock sample numbers collected by us Numbers prefixed hy 'T
Tefer to rocks collected by Professor J. L: Talbut.
242
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
77.
78.
B. DAILY and A. R. MILNES
Post-S; andalusite (A) as overgrowths on syn-S; andalusite poikiloblast, Groundmass Sj
schistosity is defined both by micas and by opaque mineral laths. Note sharp contacts be-
tween andalusite “fingers” and groundmass micas. Thin section of Petrel Cove Formation
andalusite schist SC102. Transmitted plane polarised light. Bar scale represents 0,10 mm.
Post-Si andalusite poikiloblast (A) which has crystallised around altered cordierite atugen
(C). Thin section of Petrel Cove Formation andalusite-cordierite schist T135A. Transmitted
plane polarised light. Bar scale represents 0.50 mm.
Field in Fig. 73. Crossed polars,
Post-S; chlorite poikiloblasts (c) which have crystallised across 5; and inherited the S;
fabric, indicated by elongate quartz and opaque mineral inclusions. Thin section of Tapa-
nappa Formation porphyroblastic phyllite SCS50, from ancient cliffline, western end of
Tunkalilla Beach. Transmitted plane polarised light, Bar scale represents 0.10 mm.
Post-S; chlorite poikiloblasts (c) which have crystallised across S, and inherited the S;
fabric, indicated by opaque mineral] laths. Thin section of Petrel Cove Formation andalusite
schist SC102. Transmitted plane polarised light. Bar scale represents 0.50) mm.
Post-S; euhedral garnet (g) and chlorite (c) porphyroblasts adjacent to pre-S; quartz-rich
augen (Q) in thin section of Tunkalilla Formation porphyroblastic phyllite SC58A, eastern
side of Tunkalilla Creek. Transmitted plane polarised light. Bar scale represents 0.10 mm.
Thin section of Petrel Cove Formation andalusite schist SC103 showing the development of
S» strain-slip zones, outlined by opaques. The strain-slip zones cut across post-S; chlorite
porphyroblasts {c). Note andalusite porphyroblast (A). Transmitted plane polarised light.
Bar scale represents 0.50 mm.
244 B. DAILY and A. R. MILNES
GEOLOGY OF TYPE SECTION, KANMANTOO GROUP (CAMBRIAN) 245
246 B., DAILY and A. R. MILNES
GEOLOGY OF TYPE SECTION, KANMANTOO GROUP (CAMBRIAN) 247
248 B, DAILY and A. R. MILNES
GEOLOGY OF TYPE SECTION, KANMANTOO GROUP (CAMBRIAN) 249
Sg
oe
of
B. DAILY and A. R. MILNES
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GEOLOGY
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VOL. 97, PART 4 30 NOVEMBER, 1973
TRANSACTIONS OF THE
ROYAL SOCIETY
OF SOUTH AUSTRALIA
INCORPORATED
CONTENTS
Womersley, H. B. S. Further Studies on Australian Kallymeniaceae (Rhodophyta) 253
Mawson, Patricia M. Amidostomatinae (Nematoda: Trichostrongyloidea) from
Australian Marsupials and Monotremes - - - - ae 257
Mills, K. J. The Structural Geology of the Warren National Park and the
Western Portion of the Mount Crawford State Forest, South
Australia - - - - - = “ se = =” DM:
OBITUARY: THEODORE GEORGE BENTLEY OSBORN, D.Sc., M.A., F.L.S. 317
Annual Report of Council, 1972-73 - - - - - “ z = . 391
Award of the Sir Joseph Verco Medal - - - - - - - = 322
Balance Sheet - - - - - - 2 a = - S aye are)
PUBLISHED AND SOLD AT THE SOCIETY’S ROOMS
STATE LIBRARY BUILDING
NORTH TERRACE, ADELAIDE, S.A. 5000
FURTHER STUDIES ON AUSTRALIAN KALLYMENIACEAE
(RHODOPHYTA)
BY H. B. S. WOMERSLEY*
Summary
WOMERSLEY, H. B. S. (1973). -Further studies on Australian Kallymeniaceae (Rhodophyta).
Trans. R. Soc. S. Aust. 97(4), 253-256, 30 November, 1973.
Further collections since the monograph of Womersley & Norris (1971) extend the range of
Kallymenia cribrogloea into N.S.W., clarify the structure and relationships of K. rosea and
K. polycoelioides, and show that Cirrulicarpus australis and Meredithia nana are one species now
known as Cirrulicarpus nana (J.Ag.) comb. nov. Callophyllis coccinea is now referred to as
C. rangiferinus (Turner) comb. nov.
FURTHER STUDIES ON AUSTRALIAN KALLYMENIACEAE (RHODOPHYTA)
by H. B. S. WomMersLEey*
Summary
Womersiey, H. B, S. (1973).—Further studies on Australian Kallymeniaceae (Rhodophyta).
Trans. Ry Soc. S, Aust. 97(4), 253-256, 30 November, 1973.
Further collections since the monograph of Womersley & Norris (1971) extend the range
af Kallymenia eribrogleea into N.S.W., clartfy the structure and relationships of K. rasea and
RK. palycoelioides, and show that Cirrnlicarpus ynstrafiy and Meredithia nana are one species
how known as Cirrulicarpus nana (I.Ag.) comb. nov. Callephyllis coccinea is now referred to
as C. rangiferinns (Turner) comb, nav.
Introduction
Since the monograph of Womersley & Norris
(1971) on Australian Kallymeniaceae, further
SCUBA collections by S. A. Shepherd from
New South Wales and Tasmania have provided
material permitting clarification of some doubr-
ful species, and an earlier name for Callaphyllis
coccinea is evident,
1. Kallymenia cribrogloea Womersley &
Norris. 1971: 7, figs. 6-12, 78-80.
The range of this species was previously
given as from Waldegrave I., Eyre Peninsula,
5. Aust. to Port Phillip Heads, Vic, and Bruny
L,, Tas. Well developed, typical specimens have
been found in Jervis Bay, N.S.W., 18 m deep
in the southern end of the bay (Shepherd,
15.viii.1972; ADU, A42614). This extends the
range considerably, and SCUBA collections
from deep water may show that this is not an
uncommon species, though rarely found in the
drift because of its delicate nature,
2. Kallymenia rosea Womersley & Norris
1971: 9, figs. 13-18, 81, 82.
This specics was described largely on
numerous specimens of Lucas from N.S.W.
(mainly in NSW and MBL), collected before
1912; the enly other record was one specimen
from Port Stephens, N.S,W., in ADU,
The range of K, rosea can now be extended
to Jervis Bay in southern N.S.W. where it was
found 18 m deep in the southern end of the
bay, in sheltered situations (Shepherd, 15 viii.
1972; ADU, A42615).
This material (4 specimens) agrees well in
form with the Lucas plants but is without sut-
face proliferations and is medium red in colour.
Tr is slightly rubbery when fresh und fairly thin.
The thallus of liquid preserved specimens is
(150-) 250-300 um thick (rather more than
previously estimated from dried material) and
has ao epidermis of fairly compact cells prad-
ing rapidly to large, thin walled, ovoid, inner
cells with large intercellular spaces (especially
in the centre of the thallus); fairly loose,
moderately coarse, filaments are produced from
the larger cells (Fig. 1), Some of the large cells
become stellate, as previously figured ( Womers-
ley & Norris 1971, fig. 14).
Carpogonial branch systems ate polycar-
pogonial with 8-16 carpogonial branches, the
first cell of which is clavate to lobed, and the
second cell smaller and elongate; auxiliary cell
systems dre as previously described and figured,
but the large, ovoid, cell recorded as associated
with the carpogonial branch systems was not
apparent in the Jervis Bay material. Apparently
this cell is only a yegetative cell appearing
somewhat distinctive in the Lucas specimens.
Apart from young fusion cells, stages of carpo-
sporophyte development. were not present in
the Jervis Bay matcrial.
K. resea is probably a fairly deep water
species and is distincr from other Australian
species of Kallymenia in its farm, and from
* Depurtment of Botany, University of Adelaide Adelaide, $. Aust. 5001.
254 H. B. S. WOMERSLEY
oo’ 9089 Le SANT: 090 oyeceoason0gs ee?
eo ge Oe aes ASO
OO DSSDBNDO
100.um 7
Figg 1,2,3,4
Be
Kise @H00G0 09000 08
50 um
Tes. 4a7 OD oe
AUSTRALIAN KALLYMENIACKAE
other similar foliose species in the presence of
polycarpogonial reproductive systems.
3. Kallymenia polycoelioides J, Agardh
1576) 687,
Meredithia palycoelioides (J.Ag.) J. Agardh
1892: 76; Womersley & Norris 1971: 42,
fig. 108.
Thallus complanate, foliose-subdichotomous,
ia 12 cnr high, ansing from a short stipe
1—+ mm long with a discoid holdfast 1-3 mm
across; thallus cuncately to broadly expanded
to 1-4 cm across at a few em above the stipe.
developing subdichotomous branches 1-14 cm
broad, and 1-2 cm between dichotomies, with
braad, rounded apices or lobes 3-6 mm across!
in some plants new subdichotomous fobes arise
proliferously from older basal parts. Colour
deep red to brownish-red, substance fairly soft,
adhering closely to paper on drying,
Thallus 400-600 pm thick (Fig. 23, with a
cortex 5-7 cells thick and a broad medulla of
fairly loosely arranged filaments mestly 7-15
utvin diam.; outer cortical cells fairly compact,
24 ym across and isodiametric in surface
view, inner cortical cells large: medulla with
stellate cells (Fig. 3) often with numerous arms
and slightly to moderately staining.
Curpogonial branch systems (Figs, 4, 5)
Monocarpegonial, the supporting cell lobed and
bearmg usually 4 lobed subsidiary cells and a
3 celled carpogonial branch, the first cell of
which ts lobed and the second elongate, Fusion
cell lobed (Fig. 4), Auxiliary cell systems (Fig.
7) relatively smal!, 25-50 ,.m across, consisting
of an auxiliary cell bearing about 8 subspherical
subsidiary cells each of which often bears one
further cell, Cystocarps scattered, about 2 mm
im diam. (in the type).
Male and tetrasporangial thalll unknown.
Type Locality—Orford, Tas. (Meredith).
Type—Herb. Agardh, LD (24843).
Disiribution,—South-eastern Tasmania. As
well as the lype, two fttrher Tasmanian
collections are known—Fluted Cape, Bruny
1 230 m (Shepherd, 121.1972; ADU,
A41925). and Great Taylor Bay, Bruny 1.
255
Previously this species was considered of un-
certain status {Wormersley & Norris 1971, p.
42) bur two collections by S.A. Shepherd from
near the type locality, and agreeing very well
in form with the type, show typical kally-
meniaceous female reproductive systems.
K. polycoelioides differs from other Austra-
lian species of the genus (see Womersley &
Nomis 1971, p. 4) in its subdichotamous habit,
and is most closely related to K. reba Womers-
ley & Norris in its structure and reproduction.
K. polyedelioides agrees well with Kally-
menia in thallus structure and in teproduction,
but differs from most other species in being
broadly subdichotomous (as in the holotype
illusteated by Womersley & Norris 1971, fiz,
118} rather than foliose or foliose-lobed: How-
ever, Codomier (1971) has referred two
Mediterranean species, which show 4 branched
habit. 10 Kallymenia, though without know-
ledge of the reproduction of either; K. pareny
{J.Ag.) Codomier becomes subdichoramous
several times, while K. spathulata ().Ag.)
Codomier is broadly furcate and lobed above,
K. polycoelioides has a fairly soft thallus,
drying thin and closely adherent to paper. and
not ¢arlilaginous, The texture, structure and
reproduction agree so well with other species of
Kallymeniga that K. polycoelloides cannot be
separated generically simply on the basis of its
habit,
Placement of K. potyevetioides in Kully-
menia involves consideration of the diflerence
between Aallynienia and Cirrulicarpus. The
latter is usually distinguished by its branched
thallus, Both the type species of Cirrulfcarpus
[C. gntelini (Grunow) Tokida & Masaki] from
Japan, and C. anstralis Womersley & Norris
(sce below) from southern Australia, have
thalli with several subdi- to polychotomous
branchings. While K. polycoelivides and Cirru-
licarpus both have a branched habit, the thallus
texture of the former is soft and not cartila-
ginous (as is typicul of most species of Kally-
menia), in contrast to the thallus of Cirruli-
carpus Which is cartilaginous, not or only
slightly adherent to paper, A possible distinc-
uon also lies in the cystocarps which Norris
25 m (Shepherd, 14411972; ADU,
A42131).
Fig 1. Kallymenia rosea. Cross section of thallus.
Figs, 2-7. Kallymenia polycoelioides. Fig. 2.—Cross sections of thallus with a stellate cel) and auxiliary
cel] system. Fig. 3.—Stellate cell. Fig. 4—Young carpogonia! branch system. Fig. 5—
Mature ¢arpogonial branch syslem, Fie. 6.—Fusion cell. Fig, 7, —Auxiliary cell system.
Abbreviations used: aux,c, auxiliary cell: sLe, stellate cell; subs.c, subsidiary cell; tr, trichogyne. f, 2, 3
refer to the first cell, second cell, and carpogonium of the carpogonial branch respectively.
236
et al, (1960) report as being compound or can-
fluent in €, pmelini, and which are grouped in
C. auxtrdis. For the present Cirrudicarpus is
best maintained as a distinct genus, but further
studies of distinctions between it and Kally-
menia are needed.
4. Cirrulicarpus australis and Meredithia
nana.
Womuersley & Norris (1971, p. 42) regarded
Mereatithia nana J. Agardh as of doubtful affi-
nity, commenting that the thallus structure is
kallymenivid but reproductive systems were not
geen adequately in the type. It was considered
that it might nol be a member of the Kally-
Mmeniaceac,
Further study of plants of Cirrulicarpus aus-
tralis Womersiley & Norris, especially of small
thalli, show that the type specimen of Mere-
dithia nana is almost certainly a young plant
of C. australis, Their form is comparable tak-
itig regard of the slate of development, the
structure in cross sectional view is identical.
and the type locality of M. nana (Port Phillip,
Vic.) is similar to that of C_ australis (Port
Phillip Heads).
The species should therefore be known as
Cirrulicurpus nana (J. Agardh) comb. nov,
{Basionym Meredithia nana J, Agardh 1892,
p. 76), with C. australis Womersley & Norris
(1971, p. 19, fizs, 39-43, 90) as a synonym.
H, B.S, WOMERSLEY
5. An earlicr name for Callophyilis coccinea
Harvey.
Since the publication of Womersley.& Norris
(1971), the lype specimen of Fucits rangi-
ferinus Turner has been examined. This speci-
men in the British Museum, from Kents
Islands, Bass Strait, was collected hy R. Brown
(No, 256) in 1803-4, und is a tetrasporangial
specimen 4-6 cm high, much branched with
slender ultimate branches typical of Callo-
phivllis coecinea Harvey,
‘The carrect name is thus Callophyilis rangi-
ferinus (Turner) comb, noy, (Basionym Fucus
rangiferinus Turner 1811, p, 114, pl. (83).
Following the original description of Turner,
F. remgiferinus was placed under Chondria by
C. Agardh (1823, p, 359) and under Aypnea?
by Greville (1830, p. hx) and Kuetzing (1849,
p. 761). J. Agardh (1852, p. 636) referred it
to Lecithites, but later (1876, p. 572) referred
his 1851 deseription to Mychodea harmuta
Harvey, excluding the previous synonyms. De
Toni (1897, p. 264). followed J. Agardh.
Acknowledgments
Loan of the type of F. rangiferinuy from the
Department of Botany of the British Museum
is gratefully acknowledged. Technical assist-
ance is acknowledged under a grant from the
Australian Research Grants Committee. | am
grateful to Professor R. E, Norris for com-
ments on the manuscript,
References
AuakoH, ©, (1823),—“Species Algarum"™. Vol. 1,
Pt 2, pp. 169-531,
Aqcaron, J. G. (1852).—"Species, Genera et Or-
dines Algarum". Vol. 2, Pt. 2. pp. 337-720.
(Lund).
Acaron, J. G. (1876).—"Species, Genera et Or-
dines Algarum”. Val. 3, Pt. 1, pp. 1-724.
Fpicrisis systematis Floridearum. (Lund, )-.
Acarpe, J. G. (1892).—Analecta algologiea. deta
Univ, lund. 28, 1=182, Plates 1-3.
Copomier, L,, (1971).—Recherches sur les Kally-
meénia (Cryptonemiales, Kallymeniacees), 1,
Les especes Mediterraneennes. Vie Milien 22,
ser. A. 1-54,
De Torr G. B, (1897),—‘“Sylloge Algarum om-
nium hucusque Cognitarium,” Vol 4
Florideae Sect. 1, pp. 1-388 (Padua.).
Grevitis, R. K- (1830) —"Algae Britannicae.”
(Edinburgh. ).
Kurrzinc, F. T,
(Leipzig, )-
Norns, R. E.. Toxma, J., & Masaxr, T. (1960) —
Fucther studies on Cirrulicarpus gmelini
(Grunow) Tokida et Masaki, Bull. Fac, Bish,
Hokkaido Univ. 11, 29-36,
Torver, D. (9811).—"Fuci sive Plantarum
Fucoram Generi a Botanicis Aseriptarum
Teones Deseriptiones et Historia.” Vol, 3, pp.
1-148, Plates 135-96,
Womenscey. H. B. S., & Norris, R. B. (1971).—
The morphology and taxonomy of Australian
Kallymeniaceae (Rhodophyta), Aust, J. Rot.
Suppl. Ser. No. 2, 1-f2
(1849),.—"Species Algarum,"
AMIDOSTOMATINAE (NEMATODA: TRICHOSTRONGYLOIDEA) FROM
AUSTRALIAN MARSUPIALS AND MONOTREMES
BY PATRICIA M. MAWSON*
Summary
MAWSON, PATRICIA. M. (1973). -Amidostomatinae (Nematoda: Trichostrongyloidea) from
Australian marsupials and monotremes. Trans. R. Soc. S. Aust. 97(4), 257-279, 30 November, 1973.
This work is a revision of the genera Austrostrongylus Chandler and Nicollina Baylis. Two new
genera are proposed, Paraustrostrongylus and Woolleya; these four genera, with Patricialina Inglis,
belong to the Amidostomatinae. Filarinema Ménnig is transferred to the subfamily
Mackerrostrongylinae. Austrostrongylus and Paraustrostrongylus spp. are recorded from a
phalanger and from macropod marsupials, Nicollina from monotremes and the numbat (a dasyurid),
and Woolleya from dasyurids and a native eutherian, the water rat.
New species described are Austrostrongylus hypsiprymnodontis from Hypsiprymnodon moschatus;
A. paratypicus from Macropus rufogriseus; A. chandleri from Macropus bicolor and
M. rufogriseus; Paraustrostrongylus bettongia from Bettongia cuniculus; P. trichosuri from
Trichosurus vulpecula; Nicollina calabyi and N. inglisi from Myrmecobius fasciatus; Woolleya
sprenti from Dasyurus viverrinus, Antechinus stuartii, Dasyurops maculatus and Thylacinus
cynocephalus; W. hickmani and W. monodelphis from Antechinus stuartii; W. martini from
Antechinomys spenceri.
Species redescribed in whole or in part are Austrostrongylus macropodis Chandler, A. wallabiae
Johnston & Mawson, A. aggregatus Johnston & Mawson, A. minutus Johnston & Mawson,
A. thylogale Johnson & Mawson, Paraustrostrongylus potoroo (Johnston & Mawson) (syn.
A. potoroo), and Nicollina echidnae Baylis.
Other new combinations are Woolleya sarcophili (Cameron), W. cathiae (Inglis), W. iota
(Mawson), W. acinocerus (Mawson), all transferred from Nicollina, and W. hydromyos (Mawson),
transferred from Austrostrongylus.
AMIDOSTOMATINAE (NEMATODA: TRICHOSTRONGYLOIDEA) FROM
AUSTRALIAN MARSUPIALS AND MONOTREMES
by Parricia M. Mawson*®
Summary
Mawson, Parricta M. (1973)—Amidostomatinae (Nematoda : Trichostrongyloidea) from
Australian marsupials and monotremes. Trans. R. Soc. S. Aust, 97(4), 257-279, 30
November, 1973.
This work is a revision of the genera Austrostrongyluy Chandler and Nicollina Baylis. Two
mew genera are proposed, Puraistrestrongylus and Woolleya; these four genera, with Palricia-
lina Trelis, belong to the Amidostomatinae. Filarinema Ménnigz is transferred to the subfamily
Mackerrostrongylinae. Anstrostrongylas and Parcustrostrongyluy spp, are recorded ftom a
phalanger and from muacropod marsupials, Nicollina from monoiremes and the numbat (4
dasyurid), and Wwalleya from dasyurids and 4 native eutherian, the water rat,
New species described are Austrostrangylus hypsiprymnodontis from Hypsiprymnedon
moschaius; A, paratypicus from Macrapuy riefogriseus; A. chandleri from Macropus hicelar
and M, rufogriseus; Paraustrostrongylys bettongia from Bettongia cunienlus; P, trichosuri trom
Trickosurus vulpecula; Nicollina calabyi and N- inglisit from Myrmecohius fasciatus; Woolleya
sprenti from Dasyurus viverrinus, Antechinus stuartii, Dasyurops maculatus and Thylacinus
epnocephalis: W, hickmani and W. monodelphis from Antechinus stuartii; W. martini from
Antechinomys spenceri.
Species redescribed in whole or in part are Austrostrangylus macropodis Chandler, A.
wallabiae Johnston & Mawson, A. averegatus Johnston & Mawson, A. minutus Johnston &
Mawson, A. thylogale Johnson & Mawson, Peraustrostrongylus potoroo (Johnston & Mawson)
(syn. A. poloreo), and Nicollina echidnae Baylis.
Other new combinations are Woolleya sarcophilé (Cameron), W. cathiae (Inglis), W. iota
(Mawson), W. déinocertts. (Mawson), all transferred from Nicollina, and W. hvdramyos
(Mawson }, transferred from 4 ustrostrengylus.
Introduction
The trichostrongyloid nematodes from Aus-
tralian vertebrates were discussed by Inglis
(1966) who considered that all belong to the
family Amidostomatidae. In the subfamily
Amidostomatinae he included (from marsu-
pials) the general Filarinerma MGnnig, Patricia-
lina Inglis, Anstrostrongylus Chandler and
Nicollina Baylis. Filarinema, however, differs
markedly from all other genera of the sub-
family in the virtual absence of a buccal cap-
stile—the teeth occur in an enlargement of the
anterior end of the oesophagus, so this genus
should be referred to the subfamily Mackerra-
strongylinae Inglis, in which a buccal capsule
is absent. Of the three genera from mono-
tremes and marsupials remaining in Amido-
stomutinue, no Parrictalina sp, has been found
during the present study,
Up to the present, most of the species from
marsupials have been identified as belonging
to either Austrostrongylus (type species A.
macropedis Chandler, 1924) or WNicollina
Baylis, 1931 (type species N. tachyglossae
Baylis, 1930), Proposing the genus Nicollia
(which he changed to Nicollina in 1931)
Baylis stated that this genus differs from
Austrostrongylus in the presence of a shallower
buccal capsule, the symmetrical bursa, the ab-
sence of ventral teeth, and in the shape of the
tail of the female. More species were later attri-
buted to each genus, and Mawson (1960, p.
264) pointed out that among species with ven-
tral teeth, the female tail in some: is conical
and in others bears a terminal spine, and it was
suggested that the best character on which to
separate the gcncra would be the shape of the
spicules, which in Avstrosirongylus spp. are
* Zoology Department, University of Adelaide, Adelaide, S. Aust. 5000.
15%
entire and in Nicollina spp. are bilid or trifid
distally, Inghs (1968, p. 336) agreed with this
and pave an amended list of species belonging
to each venus.
In the present study, 13 new species of ani-
dostomes haye been identified from maono-
wemes und from 4 variely of Australian mar-
supials, inzioding, dasyurids, from whith only
oie species had previously been described (N.
caihiae Jaglis, 1968). The identification of
these hus entailed the re-examination of all
available puratype or holotype specimens of
existing Species, and uw re-evaluation of the
generic characters.
Discussion
Two qew genera are proposed, Paraurtro-
iromgylus nd Woeolleva. Both Parausiro-
strongylus spp. aid Ausrrostronyyhey spp. are
characterised by wide and thick lateral alae and
and single-tipped spicules, and these features
distinguish them from Nicelltna spp. and
Woolleva spp. in which lateral alae if present
are small aod thin and the spicule tips are
divided, Distinct differences in the form of the
buceal capsule and dorsal looth distinguish
Nicollina app, trom Woalleya spp, These tour
genera are found in different hosts, A itra-
srrongvlus spp. and Paranstrosivengylus spp.
‘being recorded fram macropods and phalangers
(herbivores), Nicellina spp. from the echidna
und the numbat (termite and anteaters} and
the platypus. (which eats worms, etc.) and
Woolleya spp, Crom dasyurids, bandicoots, and
a eutherian, the water rat Call camivores),
However, there are some species which are
not satisfactorily accounted for Lhe spicules
of A. cameroni (Mawson), W. sarcophilt
(Cameron) and W, acinocercus (Mawson) are
not divided, but the body lacks the lateral alae
utd the characteristic bursa of Anstro-
virenoryins and Paruusirestrongylus,. In Waol-
leva nnvnodelphiy musp. and BO hydromyes
(Mawson) the spicules are divided and the
loneitiidinal crests are like those of Woolleva
spp. hut the tail of the female is pointed and
lacks a spine. The position of these species will
be discussed under Woulleya nig.
‘The characters of the four venera will be
described in detail below, but some general
remarks are made here about the cuticular
swellinis and crests commonly found in species
of all these genera. Cuticular ridges and cuti-
cular inflations in the Heligmosamatidae have
been studied in detail by Durette-Desset (1964,
|966) in species {rom the Old and New
Worlds, According to this author almost all the
PATRICIA M. MAWSON
heligmasarnes are rolled into a sinistral spiral.
Some live coiled around villi (with the anterior
end wwards the base of the villus) und main
tain their position by a combination of pinch-
inz with the internal (best devcloped) crests
uod the apposition of lateral crests and glae
Others, in which different arvas of the body
beur the best-develuped crests. also form a
sinistral coil, but not around a villus, and these
move through the mucus between the villi with
« corkscrew action, so |hat the outside of the
coil (the dorsal body surface) which bears the
best-leveloped crests, comes in contact with
the villi.
Although they belong to a different group
from the Heligmosomatidae, the Australian
amidostomes muy be compared with them itt
yome respects, If they wre cuiled, it is in a
sinistral spiral. Im some genera 4 4¢atro-
strongelus and Paraustrostrongylus) the Sateral
afag are Very well developed, not only out from
ihe body in a lateral direction, bul they are
wlmost 2s thick, dorsoventrally. as the body
(Fivs. 7, 13, 26). The cuticle over these en-
larged alac bears one or more Jongitudimal
ridyves, and there are alsa one of morc crests
on the ventral surface of the body proper,
These alae differ from those described by
Durelte-Desset as enlarged crests, because the
enlargement is caused by subeuticular swelling,
the crests themselves being of normal size (as
indicated by their cuticular “skeleton") and
merely borne on the inflated cuticle. In other
genera, Nicollina and Woolleya, lateral alae are
less distinct, and the loneitudinal crests are
variously distributed, In all cases males and
females of the sume species show similar pecu-
liarities of structure and distribution of the
erests. In some species, but hy no means in all,
there is 4 Bradient "round the bady jn the size
of the crests. In many Australian species, crests
are ohbsent on the dorsal surface. An attempt
has heen made to classify the various specics
according to alae and crests. Those with twa,
wide, thick lateral alae form a natural group
(also distinguished by other characters) which
is further differentiated into the two genera
Austrostrongylns and ~~ Paraustrostronyylus.
Among the remaining spectes, in which lateral
alae if present are of a different type, longi-
tudinal crests are variously arranged-- they may
be many (8-20 of more) and distributed all
round the body, or they may be few (2-4), and
restricted to the ventral surface. However,
acither of these types 38 associated with any one
of the lypes of spicule tips, dorsal ray or female
AMIDOSTOMATINAR FROM AUSTRALIAN MAMMALS 239
tail, so that further classification op the type
of crests is impracticable.
Most of the specimens cxamined were nol
seen jn situ in the host, although a few were
found coiled round torn off villi,
Unless otherwise jidicated, all the trans-
verse sections figured Were taken about the
rudhody of the worm
Measurements of new material are given in
Tables 1, 2, 3, or 4, as indicated in each case
iinder the species heading. Types of new spe-
cies Will be deposited in the South Australian
Museum, Adelaide.
Key to genera
|. Latera| alac wide and thick; dorsal Jobe of
hursa markedly thickened 2
1. Lateral alpe, if present, thin; ‘dorsal Jobe of
bursa fot thickened
2. Genital cone well developed, ‘hitinised
Paranstrosironyy lias
2. Genital cone not well developed, not chiti-
hised Alistrastrongylus
3. Buces! capsule a shallow ting; dorsal ioath
hhunt, protrusible ., _.. Nivallina
Buccal capsule # domed anteriorly: darsal tooth
pointed Weolleya
AUSTROSTPRONGYLUS Chandler
Amidustomatidac: Small usually coiled worms
with thick «nd wide Jateral alge: Jonzitudinal
cuficuJar crests on alae aml on body proper:
cephalic cuticle inflated; buccal capsule well
developed, with one dorsal and sometimes twa
smaller subventral oesophageal teeth. Male:
bursa more or less symmetrical, lateral lobes
long, dorsal short and thick; dorsal ray usually
dividing into three pairs of branches; externu-
dorsal ray arising separately, other rays from
same root, separating al tips; genital cone not
distinct, spicules slender with simple points,
sometimes united by small alae. Pemale: tail
tapering to long point; vulva heur posterior end
of body, ovejectors divergent, Parasites of
small intestine of macropod marsupials,
Type Species: od, macropodis Chandler,
1924: 160,
Other species: A. ageregatus Johnston &
Mawson, 12406: 472; 4A. minutes Johnston
& Mawson, 1938; 195; 4. rhylogale John-
stan & Mawson, 1939: 534; 4, paratypicus
nap., 4. chandleri nsp.; A. hypsiprymnodon
TSP.
In some species of Austrusirurgy lus, perhaps
in all, there is a strong tendency to {he deposi-
tion of a dark reticular and granular material
under the cuticle. Durette-Drssel (1966, pp.
457, 461) notes a similar conditiun in same
heligmosomes and. hy staining, concluded that
ln
sul lobe obscures the final branching
it is a chitinnid subslance. This occurs espe-
eially in the lateral alac aud if the bursa, and
sometimes in the cephalic inflation, but may
also appear in the older female between vulva
and anus, all round the bawdy. The distribytion
in the bursa appears to vary with the species;
in many cases it obscures the dorsal ray, espe-
clally in jong- stored specimens: it is resisten
to clearing in lactophenol, but less so in creo-
sote or Berlese's Fluid.
Tn all species of Austrostrongylus the dorsal
lohe of the bursa, whether containing granular
material or not, is: much thickened, so much so
that the lohe is in fact almost spherical, the
inner or ventral face of the lobe extending to
the cloaca, A genital cone, as such, is absent.
Because of the shape of the bursa, the dorsal
ray must be considered in three dimensions
Tather than two.
The characteristic bursa, and the striking
strap-like form given to the body by the wide
thick alac, appear to be important diagnostic
features of the genus, allhough they have not
previously been mentioned. Both characters
are present in the type species of the xentis,
which is partially redescribed below,
Austrostrongylus macropodis Chandler, 1924:
160, From Bennett's Kangaroo, Macropus
rufogriseus var, bennetti,
FIGS. tf, 2
The type and paratype material of A. praero-
podis fas been examined, The holotype male
and cotype female (U.S.N.M- Helm. Call.
26124) are va slides and impossible to exa-
mine thoroughly. What is assumed to be the
piralype material is in the collection of the
Department of Biology, Rice University. This
material (H,N. 23049) is. labelled ‘from Ben-
nett's Kangaroo’ and was worked on by Dr.
Chandler. The specimens agree closely in most
particulars with Chandler's description af 4.
mocropodis. They have the wide Jateral alac
and the swollen dorsal lobe of the bursa which
have been seen in all other species of the
genus. Unfortunately the darkening of the dor-
of the
dorsal ray. It is clear, however, that the dark
part of the dorsal jobe extends nearly to its
posterior border, and that my Chandlec’s Vig. 3
the clear part of the lobe is its swollen inner
surface, which is unpigmented. afd which in
some positions of the bursa appears to be its
posterior border. Thus the dorsal ray extends
nearly to the posterior edge of the bursa, and
the dorsal lobe is shorter that) indicated hy
Chandler. There may he two or three pairs of
2h
branches of the dorsal ray. but the tips of the
inner branches shown here in Fig, 1 are not
clear. If there are only two pairs, this is the
only species of the genus in which this is so.
The tips. of the spicules are enlarged by alae,
as figured by Chandfer.
The bodies of these specimens ure so much
contracted thai a good transverse section could
not be drawn, Allowing for some distortion,
the sections made show alae and crests similar
to those shown in Figs. 14 and 18, but with an
extra crest on the dorsal aspect of the right
side,
Key to species of Austrostrongytus
|. Female nvonedelphou. 4. rypeipremepdants
1. Female didelphous
2. Vagina lang, more or tess equal to distance
from vilvu lo anus A. aguregaius
2 Vagina short
Splewles with rehitively wide alae round tips 4
Spictiles with small or no alae at tips... 5
4. Male 45-5 mm Jong, spicules 375-500 «wm
A. macrapodis
4+. Male 3,8-4.7 mm long, spicules 520-700
ain . A, paratypicus
§. Dorsal ray gives off first pair ‘of branches, then
bifurcates ; A, minutes
5, Dorsal ruy gives off two pairs of branchex,4 then
bifurcates rie at
6. Tips of spicules united in small ala... . 7
6. Tips of spicules not united in ala
A, wallabfae
7, Three crests on each Isteral ala; branches of
dorsal tay elongate... . A, thylagale
7. One crest on each | lateral ala; branches of
dorsal ray short, stout A. chandleri
Austrostrangylus wallabiae Johnston & Maw-
son, 1939: 534, from Macropus mufrogriseus
(syn. M. ruficollis).
FIGS. 3-6; TABLE i
Host and locality: Macropus rufogrisens from
Logan Village, Qld.
New specimens a* well as the type material
have been examined and the original descrip-
tion can now be amended. As was stated. the
type specimens are darkened by masses of
granular material deposited under the cuticle
in the wide, thick, Jateral alae, in the bursa,
especially in the dorsal lobe, and in the female
al the posterior end of the body—in a few of
the older specimens this tegion is so distended
as to overhang the anus. In the newer speci-
mens from Qucunsland there are similar dark
masses biti these are neither so thick nor so
dark.
On the broad lateral alae there are longi-
tudinal crests, three on one side, two on the
other: there are wlgo two large and one small
ventral crests, There are no dorsal crests except
those on the dorsal side of the alae (Fig 1),
hed Nand
PATRICIA M. MAWSON
In the feniale the lateral alac terminate at about
the level of the vulva, in the male the left ala
reaches nearly to the bursa, and the right
rather shorter,
The buccal capsule is well developed with
two small ventral, and one large dorsal, teeth,
The spicules are very slender, with simple
acicular tips not enclosed in alae. The bursa is
thick-walled, especially the dorsal Jobe, and is
more or less symmeirical—in some specimens
the right lobe is rather longer and narrower
than the left. The dorsal ray, seen more ¢learly
in the new material, has one more pair of
branches than origitrally described (Tig, 4).
The vulva in the original material 1s 500-600
pmo. from ihe posterior end of the worm (not
1,500 pm). The cuticle just in front of the
vulva is more or Jess inflated (Fig. 2).
Although the species is similar to A. macro-
podis, there are distinct differences in the bursal
rays, and there appears to be less granular
miaterial deposited in various parts of the bady,
Austrostrungylus aggregatus Johnston & Maw-
son, 1940; 472, from Wallabia bicolor (syn.
Macropus nellahatus).
FIGS, 7-10; TABLE |
Host and locality: Wullahia bicelor
Logan Village, Old.
The type male und female and the paratype
material of this species have been re-examined
and compared with the new material. Measure-
ments of the ncw matertal are given in Table |
Two asymetrical laleral alae are present, a3
well as. three ventral longitudinal crests. The
oesophagus widens gradually in its posterior
third.
The vagina is unusually long, reaching a
distance anterior to the vulva about equal to
that of the vulva from the posterior end of
the body,
The bursa is thick, particularly the dorsal
Iohe, and darkened with 4 granular deposit. In
one male this deposit is almost absent. and
the branches of the dorsal ray quite clear (Figs,
8, 9). The spicules. are long and very slender,
hoth ending in one als. The pobernaculum is
4 thin plate.
from
Austrostrongylus minutus Johnston & Mawson,
1938: 195, from Macrepus dorsalis.
FIGS. 11-12
The paratype inaterial of this specws has
been examined; the lateral alae and the dorsal
Iohe of the bursa agree with the revited delim-
tion of these sinictures in A ustrostroniyyitey.
However, the few specimens have been greatly
Figs.
Figs,
Figs.
Figs.
Figs.
3-6,
7-10.
11-12.
13, 14.
AMIDOSTOMATINAE FROM AUSTRALIAN MAMMALS 261
i iledith Lies macropodis. Fig, 1.—Posterior end of female, Fig. 2—Posterior end
of male.
Austrostrongylus wallabiae. Fig. 3.—Transverse section of body of female. Fig, 4— Pos-
terior end of male. Fig. 5.—Dorsal ray. Fig, 6.—Posterior end of female.
Austrostrongylus aggregatus. Fig. 7,.—Transverse section of body of male. Figs. 8 and 9.—
Lateral and yentral views of dorsal ray, Fig, 10.—Posterior end of female.
Austrostrongylus minutus. Fig, 11.—Lateral view of dorsal lobe and dorsal ray. Fig,
12._Ventral view of dorsal ray.
Austrostrongylus thylogale. Fig. 13.—Transversc section of the body, Fig. 14.—Laterail
view of dorsal lobe, with dorsal and one externo-dorsal rays,
Figs, 3, 6, 7, 8, 11 and 13 to scale beside 6: Figs. 9, 12 and 14 to scale beside 12.
242
flittlened abd a useful transverse section can-
not be given. There appears to be only one
ventral hody crest, apart from those on the
alae, The lateral alue extend beyond the vulva
in the female, and in the male the left ala is a
little longer than the vight. The right side of
the bursa is rather longer than the left, but the
rays are similar on the two sides, The dorsal
lohe is so swollen as to be almost spherical,
and the three pairs of branches of the dorsal
ray (not two pairs) lic wlmost at Tight angles
to its main pxis.
Austrostrongylus thylogale Johnston & Maw-
gon, 1940a: 99, from Maeropus engenit
(syn. Thylogale eugenii), from Kangaroo 1,
5. Aust.; Mawson, 1959: 155, from Setonix
hrachyura, trom Rottoest I, W, Aust.; Inglis,
19OR! 336, from S. brachvura. W. Aust.
PIGS. 13-14
Specimens from the Kangaroo I. wallaby
have becit studied and compared with those of
other species ol Austrostrongy/us. The Jateral
alae wre broad and wide, asyoumetrical in sec-
tion, With three asymmetrical crests on the alac
and Wo Ventral crests on the body.
A. thylogale appears to be free from: the
granular Ucposits which obscure, or partially
obscure, the bursa uf some other species of the
genus, The swollen dorsal lobe of the bursa is
clear, and in it can be seen the three pairs of
branches of the dorsal ray, penetrating the lobe
in three planes (Fig. 14). In the female the
{aternl alae extend to just behind the. vulva; in
the male the left continues nearly to the bursa,
the right ends shortly anterior to this. The. spi-
cule fips are united in a very small ala.
Anstrosirongylys paratypicus 0.sp.
Fas. 15-18. TABLE |
Host anu localitys Marropus rufogriseus from
the Bathurst district. N.S.W
In the same host animal us specimens of
Austrestrangylus wullabiae, there were ubout
20 specimens of a smaller and apparently new
Austrostrongylis sp. The body form is similar
to that of 4. wallabiae, more or less tightly
cotlecL The Jateral alae are very wide, wilh one
crest on the lett side and two on the right, and
there ure in addition two ventral crests. ‘To-
wards the posterior end of the male the Icft
ala terminates. but the right, in most specimens
dark with granular material, continues nearly
to the fursa. In the female, the lateral alue
extend to or a little beyond the vulva.
The buccal capsule is well developed, the
dorsal tooth larze and the two sub-ventral teeth
PATRICTA M. MAWSON
small, The nerve ring surrounds the oesophagus
towards the end of the second third of its
length. The excretory pore is close to, oF
behind, the buse of the oesophagus in the mele,
rather More anterior in the female.
The tail of the female tapers to a Jong cylin-
urical process, Hoth ovejectors are well deve
loped, A ulerine egg near the ovejector m& 90 x
45 pm.
The bursa is thickened with granular mate-
Mul, especially in the dorsal lobe, where 2 offen
obscures the detail of the dorsal ray. The
lateral lobes. are asymmetrical, the let wider
than the nght. The three lateral aud twa ven-
tral rays are closer together in the right Yobe,
diverging only near their tips, The eaxterno-
laleral ray of the left side is distinctly larger
than that of the tight. The dorsal ray gives off
two branches before its finul bifurcation, Tot
all in the same plane (Fig. 18),
‘Vhe spicules widen near their distal ends; the
tips are wate, the alae are folded around the
lips when lying in the body and when dissesteu
out; in no case were the spicules extruded
naturally. The gubernaculum is very small andl
thin.
The species is distinguished from: A. walla-
biwe by the more posterior position of the
vulva, the shape of the spicult tips, and the
asymmetrical bursa. It seerns to be close to 4.
macrapadis (from the Vasmanian sub-species
ol Macrepas mafogrisexs). However, the spi-
cules are longer, Because of this and the appa-
reni difference in the branching of the dorsal
fay, and also because the hosts come from
widely different localities, the two species are
regarded as separate.
Austrostronpylus chandleri n.sp,
FIGS. 19-24; TABLE |
Hosts and localities: Macropus rufogrisens
(type host) from Waedlabia bicolar Tram
Lovun Village, Qld.
‘The body is loosely coiled. The lateral asym
metrical alae are well deycloped, each with one
crest; in addition, Iwo ventral crests ure pre-
sent. The buccal capsule is well developed, the
dorsal tooth about half the depth of the cupsule
und the two sub-ventral tecth smull.
The tail of the femule tapers, ending in a
thin finger-like piece. The vulva, Well in front
of the anus, leads to 4 short vagina; the ove-
jectors are about equal in sive. The cges are
75-R5 x 45-50 pm,
The spicules are long, slender throughout
their length, and end in blunt tips siightly
curved ventrally and.enclosed in a single small
263
AMIDOSTOMATINAE FROM AUSTRALIAN MAMMALS
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Figs. 15-18.
PATRICIA M. MAWSON
304m
Austrostrongylus paratypicus. Fig. 15.—Head. Fig. 16-- —Transverse section of body of
male. Fig, 17.—Posterior end of female. Fig. 18.—Bursa of mule.
Austrostrongylus chandleri. Fig- 19.—lead, Fig. 20.—Transverse section of body of male.
Fig. 21.—Posterior end of female. Fig. 22.—Lateral view of bursa. Fig. 23.—Ventral
view of dorsal ray. Fig. 24—Tip of one spicule.
Fies, 15, 19 and 20 to scale beside 20; Figs. 16, 18, 22 to scale beside I8; Figs, 23 and 24 to scale
Figs. 19-24.
beside 23.
oval ala. The gubernaculum is small and in-
conspicuous,
The bursa is more or less symmetrical, the
dorsal lobe thick, the genital cone absent as
such, The granular thickening of the bursa is
strongly developed, darkening much of it, in-
cluding the dorsal lobe and the area posterior
and dorsal to the externo-dorsal rays.
The species resembles 4, wallabiae in the
arrangement of the bursal rays (Figs, 22, 23).
It differs however in the distribution of the
granular material in the bursa, in the absence
of a lateral ala extending nearly to the bursa,
and in the shape of the tips of the spicules.
The form of the bursa and shape of the spicule
tips are close to those of A, thylogale, but in
AMIDOSTOMATINAE FROM AUSTRALIAN MAMMALS
this species the bursa is without granular inelu-
sions, the spicules are shorter, and the shape of
the dorsal ray is different.
it will be seen from the measurements given
in Table | that the specimens from H’. bicolor
are sinaller that: those from the type host.
There is also some difference in the position of
the longitudinal crests on the lateral alae in
gpecimens from the two hosts, although this is
similar in males and females for the same host,
and in collections from the same host species.
The value of this difference is uncertain; no
other morphological difference can be seen
heween the specimens.
Austrostrongylus hypsiprymnodantis n sp,
FIGS. 25-29; TABLE 1
Host and locality: Aypsiprymnodon moschatet
from Qtd,
This is.a small coiled worm, with very well
developed lateral alae, af which the right is
wider than the left. There are five longitudinal
crests, two on each Jatetal ala and one on the
ventral surface of the body. In the female the
right ala is a little shorter than the left, which
widens considerably before ending at ahout the
level of the vulva. In the male the two alac
extend from just hehind the cephalic inflation
nearly to the bursa, where they end abruptly
ut the same level,
The buccal capsule is well developed. The
dorsal tooth is ahout half the length of the
capsule. The exeretory pore lies well behind the
oesophagus.
The bursa is symmetrical, the dorsal lobe
short and swollen and with a vesiculale struc-
ture internally, which rather obscures the dorsal
ray, The arrangement of the rays is shown in
Figs. 28, 29. The spicules are long and slender
and end in a point, The presence of tenminul
alae could not be determined. A small guber-
naculum 18 present,
The tail of the female is about half the dis-
tance [rom vulva to posterior end of the body.
Only one, the anterior, uterus and ovejector is
present, Eggs wre 63-65 x 35-37 yum.
This species has been referred to Ansrro-
strorgylas because of the inconspicuous genital
cone, the origin of the externo-dorsal ray, and
the way whe lateral alae in ihe male end at the
same level just in front of the bursa. In these
features it differs from Paraustrestrangylus spp.
It is distinguished from other species of the
genus by the monodelphous condition of the
female and the posterior position of the vulva,
The cuticular swelling around the vulva is of
265
the irregular’ form seen im other species of
Ausvostenueylus rather then the definite ven-
tral owerowth seen in Parairtrostrangylur spp-
PARAUSTROSTRONGYLUS nz.
Amidostomatidae; Small, usually coiled, worms
with thick, wide lateral alae: longitudinal crests
present on alae and on rest of body; cephalic
inflation present; Ventral cuticle at posterior
cod of body inflated, forming prebursal, or pre-
vulvar, swellings usually of distinctive shape.
Buccal capsule small, with dorsal oesophageal
tooth, Ventral tecth present or absent. Male:
burs of two lateral and one small dorsal lobes,
thickened dorsally and usually ventrally; ex-
terno-dorsal ray arising from base of dorsal,
dorsal dividing into three pairs of branches,
ventro-venttal tay diverging markedly at its
base from latero-ventral and lateral rays;
genital cone strongly chilinised, spleules long
and slender, ending together in one or two alae;
guhernaculum small, plate-like; Female: tail
tapering to long point; monodelphous, only
anterior ovejector, uterus, and ovary developed_
Parasites uf the intestine of macropod mar-
supials.
Type species: P. potorac, syh. Aisitro-
strongyliy potoroa Johnston & Mawson,
1949,
This genus is close to Ausrtrostrongylus in
the form of the lateral alae, the swollen dorsal
lobe of the bursa, and the tapering tail of the
formale, It is distinguished trom it by the pre-
sence of a well developed genital cone, diver-
gent ventro-ventral rays in the bursa, and a pre-
borsal cuticular inflation in the male, and by
the absence of the posterior part of the repro-
ductive system of the female.
Paraustrostrongylus patoroo (Johnston & Maw-
son, 1969: 64), from Pororeus tridactylus.
FIGS. 30-33; TABLE 2
Host and locality: Pororous tridactylus from
Hobart; Tas,
The type material of this species, as well as
fresh material from the same hosl species, has
been examined and a fuller description is now
given,
The Jeft lateral ala is thicker than ihe nght;
each bears two crests, and in addition there are
two ventral cuticular crests and one dorso-
lateral on the right side, These crests com-
mence in the oesophageal region and continue
to about the level of the vulva in the femafe
and nearly to the bursa in the male, In the male
the Jeft ala becomes greatly inflated just an-
PATRICIA M, MAWSON
25-29. Austrostrongylus hypsiprymnodontis. Fig. 25.—Head of female. Fig. 26,—Transverse sec-
tion of body of male, Fig. 27.-—Posterior end of female. Fig. 28—Posterior end of
30-33. Purausirostrongyliy potorov. Fig. 30.—Transverse section of body of male. Fig. 31,—
Transverse section of male shortly in front of bursa. Fig. 32,—Posterior end of male.
Figs.
male, Fig. 29.—Dorsal ray,
Figs,
Fig. 33.—Posterior end of female.
Figs. 26, 27 and 28 to scale beside 28: Figs. 29, 30,
terior to the bursa: the right ala ends a little
anterior to this inflation. The subventral cuticle
on the right side anterior to the bursa is raised
into an elongate “blister” on which there are
three obliquely longitudinal crests (Fig, 31).
The buceal capsule is small and the dorsal
tooth relatively large. The ventral teeth are
very small. The nerve ring lies just behind the
mid-length of the pvesophagus and the excre-
tory pore is near its base.
The tail of the fernale tapers to a long point.
The cuticle anterior to the vulva is greatly en-
larged, forming a rounded mass overhanging
34, 32 and 33 to scale beside 32.
the vulva. The shape of this mass is similar on
all new specimens cxamined, from two hosts,
but in the older specimens il is flaltened. Only
the anterior part of the female reproductive
system is present.
The bursa is slightly asymmetrical, with the
left side, and its rays, a little larger than the
right. The thickening of the dorsal lobe is. not
very marked. The arrangement of the bursal
rays is shown in Fig, 32. Fhe spicules are
simple, undivided at the tips. A ‘small guber-
naculum is present. The genital cone is strongly
chitinised, and probably acts as an actessory
guide for the spicule.
AMIDOSTOMATINAR FROM AUSTRALIAN MAMMALS
267
TARLE 2
Measurements of Paraustrostrangylus spp, Waless otherwise indicated, meastirements ave itt pn:
a
I, petorso P. bettungia P. trichogurl
Male:
Length (mm) 2,4-2,6 4,0-4,4 4,3=5,3
QOesophagus 240-275 340-380 4350-421)
Ceph. inflation 40-48 70-80) 90-100
Antr.end—nerve ring 120-150 218-230) 250-270
—excr. pore 210-230 330-400 460-510
Spicules 250-260 380-400 380-450
Gubernaculum 4045 45-50) 35-40
Female:
Length (mm) 2,7-3,0 5.7-6.0 5.0-6.7
Oesophagus 250-290 340-41) 400-435
Ceph, inflation 45-50 80-85 80-1)
Ante. end—nerve ring 130-140 20-220 230-260
—exer, pore 200-250 280-340 440-510
Tail 90-120 105-140 120-150
Postr, end—yulva 450-200) 310-370 250-330
Paraustrostrongylus bettongia n.sp.
FIGS. 34-41; TABLE 2
Most and locality: Bettongia gaimardi from
Tas.
These are relatively large worms lying in
tight or loose coils, There are two longitudinal
crests on each lateral ala and two additional
ventral crests. Each of these ends a little in
front of the vulva in the female. Shortly in
front of the bursa.of the male, the right ala dis-
appears but the left is much enlarged. Between
the termination of the right ala and the bursa
there is an elongate subventral inflation beanng
3 to 4 oblique-longitudinal crests (Fig. 38).
In nearly every specimen the anterior end is
curved back against the rest of the body. The
cephalic inflation is about a fifth of the length
of the oesophagus. The buccal capsule is shal-
low, the dorsal tooth short. The nerve ring is
at about the middle, and the excretory pore
near the posterior end of the oesophagus.
All three fobes of the bursa are thickened.
Ventral to the chitinised genital cone there is a
cuticular thickening. which merges at each side
with the ventral part of the bursa, and which is
penetrated by the ventro-ventral ray, The
arrangement of the bursal rays is shown in
Figs. 39-41. Because of the thickening of the
dorsal lobe, the branches of the dorsal ray are
not all in the same plane, The spicules end in
alae, folded around the tips. The short guher-
naculum is finely bossed.
The vulva is about three tail lengths from
the posterior end of the body. Anterior to the
vulva the ventral cuticle is greatly inflated, and
hangs over the vulva in an elongate sausage
shaped mass, The shape of this mass is similar
in all specimens from the two host animals.
No vestige was seen of the posterior part of
the female reproductive system. Eggs are
54-55 x 32-33 pm.
The specics is distinguished from P. peforeo
by the form of the bursa, which is not asym-
metrical and is thicker-walled, by the size of
the ventral tooth, the distance of the vulva from
the anus, and the shape of the vulvar flap.
Paraustrostrongylus trichosuri n.sp.
FIGS. 42-47; TABLE 2
Host and locality: Trichosurus vulpeavla from
D’Aiguillar and Camp Mt., Old.
These worms are coiled into a fairly tight
spiral from which the anicrior and posterior
ends protrude, The lateral alae are well deve-
loped, the right with three longitudinal crests
and the left with two, and there are also three
ventral longitudinal crests. In the male the
crests and right ala disappear a short distance
in front of the bursa and ure replaced by a
sub-ventral inflation. The left ala is wider jn
ihis region and terminates close to the bursa. In
the female the alae and crests are discontinued
just in front of the yulva and there is a separ-
ate small ventral inflation just antcrior to the
vulva.
The cephalic inflation is about a fifth of the
length of the oesophagus. The buccal capsule
is shallow, and the dorsal tooth small; sub-
ventral tecth are apparently absent,
In the female the tail tapers to a fine point:
the vulva is about a tail length ta front of the
anus. Eggs are about 70 x 45 pin.
PATRICIA M. MAWSON
500m
Figs, 34-41. Paraustrostrongylay bettoneia. Fig, 34—Head of female. Fig. 35.—Transverse section of
body of male. Fig. 36.—Posterior end of female. Fig, 37.—Posterior end of mule. Fig.
38.—Transverse section of body through prebursal inflation. Fig, 39.—Ventral view of
bursa, without coverslip. Fig. 40.—Lateral view of bursa. Fig, 41,—Dorsul and externo-
dorsal rays.
Figs. 34, 35, 39, and. 40 to scale heside 40; Figs. 36 and 37 ta scale beside 37,
The margin of the bursa is entire, and the
dorsal and ventral parts are swollen. The geni-
tal cone is strongly cuticularised. The arrange-
ment of the rays is shown in Fig. 46. The
dorsal ray however is partially obscured in all
specimens examined by refractive inclusions
in the bursal wall close to the mid-dorsal line;
the ray is small, as it must be restricted to the
region of the inclusions.
The species. differs from both P. poterae and
P. betiongia in the greater development of
lateral alae in comparison with the hody dia-
meter, as well 2s in the dorsal ray and the
smaller prevulvar swelling. The bursa itself is
less swollen than that of P. bertongia and more
so than that of P. poterao.
NICOLLINA Baylis
Nicellina Baylis, 1930; 550, syn. Nicollia
Baylis, 1930, nec Nuttal, 1908, nec Krit-
schewsky, 1922.
Amidostomatinae: body with longitudinal
crests and sometimes with ofe or two Jateral
alac: anterior end with inflated cuticle; buccal
capsule shallow, stoutly built, containing a blunt
dorsal ocsophageal tooth. Bursa more or less
symmetrical, dorsal lobe absent or poorly deve-
loped; dorsal ray dividing into four branches;
externo-dorsal ray arising separately or from
base of dorsal ray; lateral and ventral rays
somewhate divergent. Spicules usually bifur-
cate or trifurcale; gubernaculum present. Fe-
AMIDOSTOMATINAER FROM AUSTRALIAN MAMMALS
Figs. 42-47, Paraustrostronyylus trichosuri, Fig. 42.—Head of male. Fig, 43.—Uransverse section of
body of female. Fig. 44.—Posterior end of female. Fig, 45.—Posterior end of mule. Fig,
46.
Bursa spread aut, inside view. Fig. 47.—Tips of spicules,
Figs. 43: and 46 to same scale, 44 and 45 to same scale, 42 and 47 to same scale.
male didelphous, vulva towards posterior end
of body; tail of female with dorsal terminal
spike and two short subveniral terminal lobes.
Parasites of monotremes and Australian mar-
supials.
Type species: N. tachyglossae (Baylis).
Other species: N, echidnae (Baylis, 1930);
N, ridei Inglis, 1969: N. camereni Mawson,
1959: N. calabyi nsp.; N. inglisi asp.; N.
baylisi n.sp.; N. mundayi asp.
Through the courtesy of Dr. W. G. Tnplis
and the British Museum (Nalural History) it
has been possible to examine the type speci-
mens of N. tachyglossae and N. echidnae. In
comparing these with other species attributed
ta Nicollina, it appears that not enough con-
sideration has been given to the shape of the
buccal capsule and the dorsal tooth. In N,
tachyglossae (Pig. 48) and N. echidnae the
buccal capsule is shallow, ring-like, and stoutly
built, and the dorsal tooth is blunt and
apparently readily protruded through the
mouth (Fig. 49). In some other species (N.
cathiae Inglis and N. sarcophili Cameron) the
buccal capsule is thinner, deeper, and some-
what domed, and the dorsal tooth ts erect and
pointed and does not seem ever to be pro-
truded through the oral opening. Moreover, in
N. tachyglossae and N. echidnae the tail of the
27)
female ends in a dorsal spine and two sub-
ventral processes, Jn species with a deeper
buccal capsule this type of tail has not been
seen.
It is concluded that species having these
characteristics in common should be grouped
in a genus for which the name Nicollinw is
availahle, For the other specics formerly in-
cluded in Nicollina a new genus, Woolleya, 1s
proposed.
Key to species of Nicollina
1. Worms more or less couled .... ..,
1. Worms not coiled a oe ae.
2. Dorsal ray with ihree pairs of branches
2, Dorsal ray with two pairs of branches ..
3. Buccal ring thick, lobed anteriorly . MV. ealabyi
3, Buccal ring not Johed, thinner and deeper _..,
N- inglisi
N. ritdei
NN, eamieront
bie Yr
4. Spicules bifid
4. Spicules simple
5, Spioules trifid —_ en
5. Spicules bifid. .., a eee 7
6, One lateral ala present N. echidnae
N, fachyglossae
N. mandeyi
N. havlist
6. Lateral alae absent .
7. Lateral alac present
7. Lateral alac absent
Nicollina echidnae Baylis
VIG, 50
Nicollina evhidnae (Baylis) Baylis, 1931,
Mawson, 1959; 154; syn. Nicollina echidnae
Baylis, 1930; 14. From Vachyglossus acu-
leatus.
The material examined in 1959 from an
echuina from Kangaroo L has now been com-
pared with the type material of the species, and
the identification confirmed. A transverse sec-
tion of the Kingaroo |. specimen is given. In
this species, and in N. tachyglossue, the excre-
tory pore, not mentioned by Baylis, is post-
oesophageal.
Nicollina cameroni Thomas, 1959; 154, from
the Echidna, Tachyglosvuy aenuleatys.
FIGS, 51-52
The paratype material of this species has
been re-examined. ‘Ihe body was described as
having two lateral alae and “some appearance
of fongiludinal banding’. Transverse sections
show a very slight widening of the cuticle later-
ally and in addition about 16 crests, most of
them lateral or ventral, The tooth is blunt, and
lies for the most part in the oesophageal fua-
nel.
This species. differs from others of the genus
in that the spicules are not divided distally, but
in view of the: similacity of other characters it
has been retained in the genus, ft differs very
markedly trom spectes of Austrostronevlus and
PATRICTA M. MAWSON
Prraustrostrongylus, in which the spicules. ace
single, in characters of the lateral alae, buccal
capsule, dorsal tooth, and bursa,
Nicollina haylisi n.sp.
FIGS. $3-59; TABLE 3
Host and locality; Tachyglossus aculeatus, Tas,
‘These are straight worms, with a cephalic
inflation, followed by numerous very low longi-
tudinal crests. Lateral alae ave absent. The
buccal capsule is very shallow, The oesophagus
is more of Jess cylindrical in its anterior two-
thirds then widens to an elongate bulb at the
posteriar end. The nerve ting surrounds it just
behind its midlength and the excretory pore
and cervical papillae are at about the same
level near its posterior end.
The lateral lobes of the bursa are very long
and are folded over each other, The externa-
dorsal ray arises from the dorsal and diverges
widely from it. The dorsal ray divides into two
bifid branches near tts distal end. The spicules
ure bifid for about half their length. The outer
branch of each ends in a barb and bears about
10 well-marked transverse ridges. in the middle
third of its length. ‘The eid of each branch is
surrounded by an ala, that on the smaller
branch much wider than that on the larger.
The gubernaculum is Stout, pitted on the sur-
face, and rather more than half the sprcule
in length.
The tail of the female is rounded and bears
a subterminal spike and two small lobes, The
vulva, a (transverse sli, ina depression of the
body wall, lics at a little less than a sixth of the
bodv length, or 20-27 tail lengths, from the
posterior end of the body, Uteri are opposed.
The eggs are 79-80 x 43-45 jam,
‘Vhe species is in many ways very like N.
tachyelossae, a straight worm with 8-10 longi-
tudinal crests and with somewhat similar, but
trifid, spicules. The two species difler however
in hody length, position of the vulva, and size
of the gubernaculum,
Nicollina munduyi nsp,
FIGS. 60-66; TABLE 3
Host and locality: Tarkyglossus aeuleatus and
Ornithorhynchus anatinus from Tas.
This is a short straight worm very simiur in
some respocts to N. echidnae. There are wo
jateral alae and about 20-22 longitudinal cuti-
cular crests, more or less: evenly distributed
around the body circumference for must of 1s
length, but. fewer towards the extremities.
The buccal capsule is short, ring-shaped; the
dorsal jooth is blunt and, at rest, hardly pro-
AMIDOSTOMATINAF FROM AUSTRALIAN MAMMALS 271
Figs. 48, 49.
Fig. 50.
Figs. 5t, 52.
Figs. 53-59,
Nicollitta tachyglossae, lateral and dorsal views of anterior end.
Nicollina echidnae, transverse section of body,
Nicollina camerani, Fig. 51.—Head. Fig, 52.—Transyerse section of body.
Nicollina baylisi. Fig. 53,—Oesophageal region. Fig. 54.—Lateral view of head. Fig,
55.—Trunsverse section of body just posterior to ocsophagus. Fig. 56—Posterior end
of male. Fig. 57.—Dorsal ray. Fig. 58,—One spicule. Fig. 59.—Tail of female.
Figs. 48, 49, and 57 to scale beside 51; Figs. 52 and 53 to same scale; Figs. 54, 57, 58, and 59 to scale
beside 59.
jects into the buccal cavity. The oesophagus is
cylindrical for most of its length, ending in a
bulb, The nerve ring lies at about half, and the
excretory pore and the small but distinct cervi-
cal papillae at three-quarters the length of the
oesophagus.
The end of the tail of the female bears a
terminal spike and two small subterminal lobes.
The vulva, at about 8-11 tail lengths in front
of the anus, lies between two rounded expan-
sions of the lateral alae. Eggs are about 75 x 40
wim,
The spicules are bifid, the outer branch of
each is longer and stouter than the other and
ends in a barbed point, the inner branch ending
simply. As only two male worms are present,
the spicules were not dissected out. Their
appearance is very similar to those of N. hay-
lis?, on which the terminal alae were not visible
until the spicules were out of the body. The
gubernaculum, at least two-thirds the length of
the spicules, is strongly built and its surface
pitted. The lateral Inbes of the bursa are not
particularly Jong. The rays are shown in Fig
63.
The species closely resembles N. baylivi, but
is distinguished by being distinctly shorter, with
lateral alae and with fewer and more promi-
nent cuticular crests, as well as by the shape of
the bursa, the absence of ridges on the spicules,
PATRICIA M. MAWSON
TABLE 3
Measurements of Nicollina baylisi, N. mundayi, N. calabyi, and N. inglisi. Unless otherwise indicated,
measurements are in jm.
$e
Species N. baylist N. mundayi N. calabyi N. inglisi
Host echidna echidna platypus numbat numbat
Male:
Length (mm) 12,8-14.1 §.2,5.3 4.6—-5.7 2,4-3,.4 2.1-3.4
Oesophagus 500-525 445, 500 400-450 330-360 210-250
Ceph. inflation 80 (3x) 75,75 70-100 50-70 60-65
Antr. end—nerve ring 210-220 215, 220 220-230 120-170 100-120
—excr. pare 360-400 275, 315 370-390 160-260 1S0-180
Spicule 240-275 270, 295 250-270 600-700 400-440
Gubernacuwlum 145-165 140, 152 140-1660 80-110 70-80
Female:
Length (mm) 24.3, 25.3 6.5-8.1 7,7-8.0 2.5-4.2 2.3-3.4
Ocsophagus 600, 520 480-510 450-500 260-400 270-335
Ceph. inflation 100, 110 75-80 80-110 60-80 55-70
Antr. end—nerve ring 300, 270 210-240 230-240 120-20) 100-115
—excr. pore 500, 550 230-305 380-400 150-345 140-220
Tail 190, 150 120-140 140-165 70-130 60-90
Postr, end—vulva 3800, 4100 1250-1400 1500-1600 350-460 200-300
64
200,um
Figs. 60-66. Nicollina mundayi, Fig. 60.—Head. Fig. 6i—Oecsophageal region. Fig. 62.—Transverse
section of body. Fig. 63—Posterior end of male, ventral view. Figs. 64 and 65
and lateral views of region of vulva. Fig. 66.—Tail of female.
Figs. 61-66 to same scale; Figs. 64 and 65 to same scale.
Ventral
AMIDOSTOMATINAE TROM AUSTRALIAN MAMMALS
and the presence of culicular Maps beside the
vulva. The measurements ate similar to those
of N. evhidnae, bul the two species are dis-
tinguished by the shape and size of the spicules
and of the dorsal altd externo-dorsal rays, and
by the presence of the well-developed Icft
lateral ala in N. eehidnae.
Nicollina ealabyi n.sp.
FIGS, 67-72, TABLE 3
Host and locality! Myrmecobius fasciatus from
W. Aust,
The body forms a loose coil, The length of
the anterior cuticular inflation is about one and
a half times the hody width just behind the
inflation, and about 4 fifth to a sixth the length
of the oesophagus. The cuticle af the rest of
the body ts ratsed into 8-9 lateral and ventral
longitudinal crests, of which the lateral are thy
best-developed,
The mouth is surrounded by six small cuti-
cular lips. The buccal capsule is stourly built,
its anterior edge six-lobed, each lobe formed
by a thickening of the wall on the outer side
of the capsule. The dorsal tooth is rounded at
the apex, and reaches to about half the depth
of the buccal capsule. The cesophagus is wiser
in the second half of its length and is sur-
tounded by the nerve ring al the end of its first
quarter; the excretory pore hes at aboul the
ettd of the third quarter
The tail of the female is rounded at the tip,
with a subterminal spine af low ventral pro-
minence. The vulva is a transverse slit, three
to four (ail lengths in front of the anus, At the
evel of the vulva the cuticle is raised snto three
tongitudinal crests, one, bilobed, to the leét of
the vulva, a frilled narrow one to the right of
the vulva, and a longer wider one to the right
of this again (Fig, 72), The sizes and arrange-
mentot these crests gre similar in all the speci-
mons available The eggs are thin-shelled,
65-70 x 35-40 pm.
The spicules of the male are bifid for the
terminal 80-100 ym, the shorter of the two
ends pointed, the longer truncated and slightly
barbed, The gtbernaculum is elongate,
rounded at the ends, und thicker in the central
pacts. The bursa has a short dorsal lobe,
slightly separated From the large latero-ventral
lobes, The rays are arranged as shown in Fig.
69,
The species is placed in the genus Nicellina
because of the form of the huccal capsule and
tooth, It differs from the species frony mono-
tremes in the form of the dorsal ray, as well as
273
in the presence of 3 cuticular crests near the
vulva.
Nicollina inglisi i.sp.
FIGS, 74-76: TABLE 3
Host and locality: Myrmecohius fasciatus from
W, Aust.
This is a rather slender loosely coiled worm,
The length of the inflated cephalic cuticle is
about twice the width of the body just behind
it. There are 8-10 longitudinal crests on the
body, (wo dorsal and two Jateral, and the rest
ventral. The buccal cavity contains a large
blunt tooth. The oesophagus widens in its pas-
terior third and the nerve ring surrounds it al
about the end of the first. third, The excretory
pore is at the end of the second third.
The tail of the female lapers to & bi-lobed
tip, with a spike arising subterminally, The
vulva is a transverse slil siluated about three to
four times the tail length in front of the anus.
The cuticle just anterior und posterior to the
vulva is slightly inflated and strongly striated,
Amphidelphous, posterior uterus short, but
containing developing cgys. Eggs are up to 100
x 50 um,
In the male. one side of the bursa is slightly
longer than the other, but the rays are similarly
disposed. The dorsal lobe is short and the
dorsal ray thin (Fig. 74), The spicules are
bifid, with one branch ‘shorter than the other,
and the four tips, two from each spicule,
appear to be enclosed in one terminal ala
The species is close to N, calabyi, which was
found in the same collections. Tr is distin-
guished by the more slender build of the body,
the relatively longer cephalis inflation, the
slightly thinner-walled and deeper buccal cap-
swe, without anterior thickening; in addition,
the guhernaculum is longer, and there is a
small hut consistent difference in the dorsal
rays.
WOOLLEYA n.g.
Amidastomatinae: Small, more or tess coiled
worms with inflated cophalic cuticle and longi
tudinal cuticular crests behind this; huocal cap-
sule well developed, with pojnted dorsal tooth.
Male; bursa more or less symmetrical, dorsal
lobe small, dorsal ray dividing into two or three
branghes, externo-dorsal ray arising from
dorsal ray or separately: fateral and ventral
rays arising together, diverging from mid-
lengths; genital cane not strongly developed.
spicules single or divided at tips; gubernaculum
well developed. Female: tip of tail rounded
with thin spike, or tapering ta a point; vulva
274 PATRICIA M, MAWSON
304m
67 69
75 |il|
2
iy 71
Figs, 67-72. Nicollina calabyi. Fig. 67—Head. Fig. 68.—Transverse section of body, Fig. 69,—
Bursa. Fig. 70. -Gubernaculum and tips of one spicule, lateral view. Fig. 71.—Tips of
both spicules, ventral view. Fig. 72—Posterior end of female.
Figs. 73-76. Nicolfinw inglisi. Fig, 73,—Anterior end, Fig. 74—Bursa. lig. 75.—Tips of spicules.
Fig. 76.—Posterior end of female.
Figs 67, 70, 71, 73, and 75 to scale beside 73; Figs. 68, 69 and 74 to scale heside 69; Figs. 72 and 76
to scale beside 76.
towards posterior cnd of worm: mono- or di- W.. hickmant msp.; W. martini nsp., W.
delphous. Parasites of intestine of Australian monodelphis n,sp,
mammals, mainly of marsupials, The genus is named in recognition of the
Type species: W. sprenti n.sp. help given by Dr. Patricia Woollcy in collecting
Other species: W. cathiae (Inglis), syn. specimens from dasyurids.
Nicollina cathiae; W. sarcophili (Cameron), Woolleya species are distinguished from
syn. Nicollirta sarcophili; W- iota (Mawson), Nicollina spp. chiefly by (he shape of the buccal
syn Nicollina tota; W. acinucercus (Mawson), capsule which is cup-like with relatively thin
syn. Austrostrongylus acinocercus; W. walls, and by the shape of the sharply pointed
hydramyos (Thomas), syn. A. Aydronpyos; dorsal tooth, originating from the anterior end
AMIDOSTOMATINAK FROM AUSTRALIAN MAMMALS 134
of the oesophagus rather than, as in Nicollina
spp., from the anterior cnd of the wall of the
lumen of the oesophagus.
Figures are given of transverse sections of
W. acinocereus and W. hydromyes, which have
been re-examined but are not redescribed here
(Figs. 71, 78).
Key to specics of Woollceya
1. ‘Pail of female tapering to & point . 2
{. Tail of female rounded at tip, with spike . 4
2. Spicules not divided, or irifid! .
Ww. sareaphill
2, Spicules bifid .. 2 3
Female didelphaus WW, hydreinyiap
. Female meonodelphous WW. monodelphi
4. Spicules noi divided W. acinocerens
4. Spleules bifid... _)
5. Longitudinal create nore or "hex ‘evenly -distri-
buled around body
§. Longitudinal crests only on ventral surface. $
6, Tip of Jongcr branch of cach spleule en-
larged . |. W, jiora
6& Neither tip of spienles enlarged . theese
7. Dorsal ray ends in three pairs of branches
W. martiti
7. Dorsal ray ends in two pairs af branches
W hickmiani
8. Narrow lateral alae present W. cotkiae
8. Lateral alae absent —.. ... HW. sprenti
Woolleya sprenti n.sp-
FIGS, 79-85; TABLE 4
Host and localities: Desyuruy viverrinus from
Icena, Tas. (type host and locality); Arie-
chinus stuarti from Mt, Tidbinbilla, A,C.T,;
Dasyurops maculans from N.S.W., Thyla-
clings cynocephalas from Tas,
These are relatively long, slender worms,
some coiled louscly, some in a tight spiral; The
inflated cephalic cuticle, about a quarter the
length of the oesophagus, is lightly striated
transversely, the cuticle on the rest of the body
is more heavily striated and thrown into three
longitudinal crests extending most of the body
length on the ventral side.
The buccal capsule is large with a dorsal
tooth just over half the depth of the capsule.
Ventral teeth were not seen. The nerve ring
is. just behind the mid-oesophagus and the ex-
crelory pore near the posterior end of the oeso-
phagus.
The posterior end of the female narrows
abruptly just in front of the anus and the tail
is digitiform with a rounded tip bearing a ter-
minal spike. The vulva is about five to eighr
tail-leneths in front of the ants. Two ovejectors
ae
aud Wteri are present, the posterior much the
shorter,
The bursa is symmetrical, its dorsal lobe
short but quite distinct. The arrangement of
the bursal rays is shown in Figs. 82 and 83.
Each spicule hifurcates at abouf a fifth its
length; one branch is slightly longer and stouter
than the other, and is curved inwards at the
tip. The gubernaculuin is long and wide, the
central part more heavily chitinised,
The specimens (two females) from Thyla-
cinus cynocephalus agree in all particulars with
the types. In view of the different host, it is
possible that they may belong to a different
species, distinguishable only by characters of
the male, As the host species is now virtually
extinet it is unlikely that a male will be found,
unless in some museum. The specimens des
cribed here were found in a museum specimen
of the host, through the enterprise of Professor
J.. F. A. Sprent,
This species mast closely resembles WN.
Aydrontyos Thomas in the shape of the spi-
cules and the arrangement of the dorsal ray-
The two species differ however in the lengths
of spicules and guhernaculum and in the shape
of the femate tail.
Woolleya martini nsp,
FIGS. 86-89; TABLE 4
Host and localily; <Antechinomys
from Sandringham, Qld.
This is a relatively small species, with 10
longitudinal cuticular crests. The cephalic in-
flation of the cuticle is aboul twice as long as
tts diameter, and between a third and # quarter
the length of the oesophagus. ‘The cuticle
around the mouth forms six distinct lips, The
dorsal tooth is about half the length of the
buveal capsule. The exerectory pare lics iat or
just behind the base of the oesophagus,
The spicules are bifid in their distal quarter:
both branches are slender, one a little more
curved and slightly longer than the other. The
arrangement of the bursal rays is shown in
Fig. 88.
The body of the female narrows just in front
of the anus and ends in a digitiform tail,
rounded at the tip and bearing a terminal spine.
Only two, sub-veutral, crests continue posterior
to the vulva.
This species js distinguished from W/, sprenr
chiefly by the number of cuticular crests, and
by the origin and shape of the oxterno-dorsal
Tay.
spencert
1Tke number of terminations of exch spicule is net stated dn the description of this species.
PATRICIA M. MAWSON
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AMIDOSTOMATINAE FROM AUSTRALIAN MAMMALS
84
504m
50m
Fig. 77. Woalleya hydromyos, transverse séction of body.
Fig. 78, Woolleya acinocercus, transverse section of body.
Figs. 79-85.
Woolleya sprenti, Fig. 79.—Anterior end, Fig, 80.—Oesophageal region. Fig, 81.—Trans-
verse section of body. Fig. 82.—Posterior end of male, Fig. 83,—Dorsal and one extertio-
dorsal rays. Fig. 84.—Spicules. Fig. §5.—Posterior end of female.
Figs. 77, 79, and 83 to scale beside 77; Figs. 78 and 84 to scale beside 84; Figs, 81 and &2 to Scale
beside 82.
Woolleya bickmani n.sp.
FIGS. 90-93; TABLE 4
Host and locality: Antechinus stuartii from
Condor Creek and Mt, Tidbinbilla, A,C,T,
These are fongish coiled worms. The cuticle
is raised into ten or twelve longitudinal crests,
extending for most of the body length, and
more or less evenly distributed around the
body, the widest gap being on the dorsal side
(Fig. 91). The buccal capsule is shallow and
the dorsal tooth short.
The bursa is more or less symmetrical, the
dorsal lobe not separated from laterals. The
arrangement of the rays is shown in Fig. 92,
The spicules bifureate at about 3/4 or 4/5
their length; each branch ends in a blunt point,
PATRICIA M, MAWSON
Wooalleya martini. Fig. 86.—Anterior end. Fig, 87.—Transverse section of body. Fig.
88.—Part of bursa. Fig. 89.—Posterior end of female,
Woolleya Aickmani. Fig. 90—Antetior end, Fig. 91—Transverse section of body hebind
oesophagus. Fig. 92,—Bursa. Fig. 93.—Posterior cnd of female.
Woolleya monodelphis. Fig. 94.—Anterior end, Fig. 95,—Transverse section of body.
Fig, 96.—Bursa, Fig. 97.—Spicules. Fig. 98—Posterior end of female.
Figs. 86, 91, 94, and 97 to scale beside 94; Figs. 87, 88. 90, 92, 96, and 98 to scale beside 88.
the longer one rather more curved at the tip.
A thin plate-like gubernaculum is present.
The posterior end of the body of the female
is slightly swollen; the tail tapers somewhat and
is rounded at the end, with a terminal spike.
Two ovejectors are well-developed. Eggs are
about 60 x 30 ym.
This species differs from W. martini in the
distribution of the cuticular crests and in the
branching of the dorsal ray (see Figs. 88, 92).
Woolleyi moenodelphis nsp.
FIGS. 94-98: TABLE 4
Host and locality: Antechinus stuartji from
Condor Creek, A:C.T.
This is a very small specics; the anterior end
of the body ends in a more or less tight. spiral:
the posterior is curved, and in the Female dis-
tinctly swollen in the region of the vulya. There
are four longitudinal ventral crests, on the an-
terior two-thirds or more of the body. The
AMIDOSTOMATINAE FROM AUSTRALIAN MAMMALS
cephalic inflation is about two-thirds the length
of the ocsophagus. The buccal capsule is large,
the dorsal tooth very small. The posterior end
o! the oesophagus was secn clearly in only one
specimen, and the nerve rig and exerttory
pore were not secn im any,
The arrangement of the bursal rays is shown
in Fig. 96. ‘The dorsal pay is unusually stout,
and arises separately. The branches of the
dorsal ray are very small, and it is possible that
the final branch shown in Fig. 96 is divided.
The spicules ate hifid for ahout one third of
their Jength. one branch of each being thicker
and slightly longer than the other. A slender
gubernaculum is present.
Fhe tail of the female is relatively Jong and
tapers to a fine point The vulva lies about one
tail length in front of the anus, between two
short and wide sub-ventral crests or flaps,
There is only one ovejector and uterus, the
antenor. No eggs were seen,
This species differs from all others referred
to Woolleya in being monodelphous, in the
wide externo-dorsal rays, and in the very small
279
size, and from most of the species in the shape
of the tail of the female.
Acknowledgements
The collections used in this sludy were made
available by a number of people, Some material
from Tasmania was sent by Dr. B. Munday
and Mr. R. Green of Launceston, and some
by Or. John Hickman of Hobart. Dr. John
Calaby (C.S.LR.0. Division of Wildlife Man-
agement) sent some nematodes from a numbat;
Dr. P. Woolley contributed most of the mate-
rial from Anfechinus stuartii. "The bodies of
Antechinomys sp,, native cats, and a numbat
were donated by my colleague, Professor P_ G,
Martin. Material from Queensland wallabies
was sent by the late Dr. M, J. Mavkerras, Pro-
fessor Sprent (Dept. of Parasitology, Univer-
sity of Queensland) very kindly let me have
the two Worms from the thylacine,
All of these people have taken considerable
trouble to send worms. or host bodies in a good
state of preservation, and [ am very grateful
to them.
Most of the work was carried out while in
receipt of a Rural Credits Research Grant,
References
Bayus, H. A. (1930) —Four new: trichostrongylid
nematodes from Queensland. Ann. Mag. nat.
Hist, 1 (6), 1-18,
Bavots. A, A. (1931)—A nomenclatural correc-
tion, Aan, Mag, nat, Hist, 10 (7), 550,
Camenon, 'T. W. M. (1931).—On a species of
trichostrongyle from the Tasmanian Devil. 7,
Helminth. 9, 153-156,
Cuanpior, A. C, (1924) —A new genus of tricha-
slrongylid worms from the kangaroo. Pera-
sitology 16, 160-163.
Durette-Desset, M. (1964),—Les systémes
Warétes cuticulaires chez les nématodes hélig-
mosomes, Etude de cing espéecs parasites do
rongeurs de la Maboké. Cahiers de La
Mahoké 2, 40-78.
Digerre-DesseT, M. (1966)—Sur deux nou-
yeaux trichostrongyles purayites du Porc-Epic
au Viet-Nam. dArnntls. Parasit. hum. comp. al,
453-466.
Ineuis, W. G. (1968).---The geographical and
evolutionary relationships of Australian tri-
chostrongyloid parasites and their hosts. J.
Linn, Sac, 47, 327-347.
Jonnston, T. H., & Mawson, P.M, (1938).—
Some nematodes from Australian marsupials.
Ree. S. Aust. Mus. 6, 187-198.
Ionwston, T. H., & Mawson, P. M. (1939).—
Stronyylate nematodes from marsupials in
New South Wales. Prec. Linn. Sac. NSW.
64, 513-536.
Toumsron, T.. H., & Mawson, P.M, (194038) —
Nematodes from South Australia. Trans. R,
Soc. S. Aust. 64, 95-100,
Jounston, T. H., & Mawson, P. M. (1940b).—
New and known nematodes from Australian
tea Proc. Linn, Soc, NSW. 65, 468-
476,
Jounsion, T. H., & Mawson, PL M, (1949).—
Some nematodes from Australian hosts to-
fether with a note on Riahditis allgeni. Trans,
R. Sov. §. Aust. 7a, 63-71.
Mawson, P, M. (1960),—Nematodes belonging to
the Trichostrongylidae, Subuluridae, Rhab-
dinsidae, and Trichuridac from bandicuats-
Aust. J. Zool. 8, 261-284.
Mawson, P. M. (1961). —Trichostrongyles from
rodents in Queensland, with comments on the
genus Longistriata (Nematoda: Heligmoso-
matidae). Aust. J. Zoal. 9, 791-826,
THomas {—Mawson), P. M, (1959).—Some
nematode purasites from Australian hasts,
Trans. &, Sov, S. Aust, 82, 151-162.
THE STRUCTURAL GEOLOGY OF THE WARREN NATIONAL PARK
AND THE WESTERN PORTION OF THE MOUNT CRAWFORD
STATE FOREST, SOUTH AUSTRALIA
BY K, J. MILLs*
Summary
MILLS, K. J. (1973) -The Structural Geology of the Warren National Park and the Western Portion
of the Mount Crawford State Forest, South Australia. Trans. R. Soc. S. Aust. 97(4), 28 1-3 15, 30
November, 1973.
A structural study of coarse-grained sillimanite-bearing mica schists and gneisses exposed on the
southern shores of the Warren Reservoir, south-cast of Williamstown, has revealed evidence for
three successive phases of strong compressive deformation, each characterised by the introduction
of new structural elements. These rocks stratigraphically overlie a thick crossbedded schistose
metasandstone unit which closely resembles the Aldgate Sandstone of the Adelaide Supergroup.
This metasandstone unit also displays evidence for three superimposed fold episodes. Coarse
grained sillimanite-bearing mica schists and gneisses underlying the metasandstone have a similar
lithological and compositional range to those above the metasandstone, but are more complexly
deformed and show indications of earlier compressive deformation events. Careful mapping of the
contact between these older schists and the metasandstone has revealed an angular unconformity in
several localities, thus re-establishing an exposure of pre-Adelaidean basement, herein named the
Warren Inlier. Rocks within the inlier have been reconstituted at high temperatures along with the
Adelaidean sequence and have structural characteristics quite unlike those of the previously
described basement inliers within the lower temperature metamorphic environments of the Mount
Lofty Ranges. Detailed mapping has established that a period of major thrust moyements took
effect during and after the second compressive deformation in the Adelaidean sequence, and that
these thrusts were later folded during the third compressive deformation. The final significant
tectonic event is recorded by the Williamstown-Meadows Fault, which brought the amphibolite
facies schists and gneisses into conjunction with biotite zone phyllites.
The fortunate combination of thrusting and updoming in subsequent folding, with later uplift on the
Williamstown-Meadows Fault in this part of the central Mount Lofty Ranges, has enabled erosion
to unveil a tectonic window, herein named the Williamstown Window, through which may be read
the structural history of the lowermost tectonic and stratigraphic leyels of the Upper Proterozoic
sedimentary pile and the underlying basement gneisses.
THE STRUCTURAL GEQLOGY OF THE WARREN NATIONAL PARK
AND THE WESTERN PORTION OF THE MOUNT CRAWFORD STATE FOREST,
SOUTH AUSTRALIA
by K. J. Mitis*
Summary
Miuus, K. .|, (1973).—The Structural Geology of the Warren National Park and the Western
Portion of the Mount Crawford State Forest, South Australia, Trans, R. Soc, 8, Ausé.
97(4), 281-315, 30 November, 1973.
A structural study of coarse-grained sillimanite-bearing mica schists and gneisses exposed
an the southern shores of the Warren Reservoir, south-east of Williamstown, has ‘evenled
evidence for three successive pluses of strong comipressive deformation, each characterised by
The introduction of new structural elements. These rocks stratigraphically overlie a thick cross~
hedded schistose tmetasandstone unit which closely resembles the Aldgate Sandstone of the
Adelaide Supergroup, This metasandstone unit also displays evidence for three superimposed
fold episodes. Coatse-grained sillimanite-bearing mica schists and gneisses underlying the meta-
sandstone have a similar lithologica] and compositional range to those above the mctasandstone,
but are more complexly deformed and show indications of earlier compressive deformation
events. Careful mapping of the contact between these older schists and the metasandstone has
revealed an angular unconformity in several localities, thus re-establishing an exposure of
pre-Adeélaidean basement, hetein named the Watten Inliet. Rocks within the inlier have been
reconstituted at. high temperatures along with the Adelsidean sequence und have structural
characteristics quite unlike those of the previously described basement inliers within the lower
temperature metamorphic environments of the Mount Lofty Ranges. Detailed mapping hus
established that 4 period of major thrust movements took effect during and after the second
compressive deformation in the Adelaidean sequence, and that these thrusts were later folded
during the third compressive deformation, ‘The final significant tectonic event is recorded
by the Williarmstuwo-Meadows Fault which brought the amphibolite facies schists and gneisses
into conjunction with biotite zone phyllites,
The fortunate combination of thrusting and updoming in subsequent folding, with later
uplift on the Williantstown-Meadows Fault in this part of the central Mount Lofty Ranges,
has enabled erosion ta unveil a tecfonic window, herein named the Williamstown Window,
through which may he read the structural history of the lowermost tectonic und stratigraphic
levels of the Upper Proterozoic sedimentary pile and the imderlying basement gneisses,
the highest peak in the area and from its
central location one can view the rugged
Introduction
This paper is concerned with the elucidation
of the structural history of a key area of
approximately 40 km? in the central Mount
Lofiy Ranges. The atea is located south-east
of Williamstown and extends south and west
of the Warren Reservoir. Much of the area is
virgin scrubland, parts of which have been
recently acquired by the National Parks Com-
mission (Warren and Hale National Parks), or
are wnder the control of the Engineering and
Water Supply Department. or form perl of
the Mount Crawford State Forest. There are
a few small grazing properties in Dead Horse
Gully (Watts Gully) and south of the reser-
vair. Lookout Tower Hill (over 525 m) forms
guilied scarp country of the Warren National
Park falling off to the west, and the gorge
of the South Para River and the Warren Re-
servoily to the north. Outcrops are frequent
and fresh on these youthfully eroded slopes,
To the south-east and south a much more
mature topography, containing remnants of
laterised erosion surfaces, exhibits only scat-
tered bedrock exposures, most of which are
deeply weathered.
The area was the scene of a goldrush in
TRRS, and this. led lo the appearance of one
of the earliest geological maps to be published
in this state (Brown & Woodward 1886). In
* Deparimicat of Geology ang Geophysics, The University of Sydney, Sydney, N.S.W. 2006.
482
describing the bedrock around Watts Gully
(See Fig. J) Brown & Woodward clearly
recopnised (wo mictamorphice sequehecs—
micaceous and chloritic and clay slates with
beds of quartzite und schistose sandstunes; and
highly metamorphosed rocks consisting of mica
schist, quiartzites, sandstones and ynvisste
rocks. The goldficlds were situated in the
highly metamorphosed sequence.
Howchin (1906, pp. 254-256), referring to
the ridge between Williamstown and the Suuth
Para River. recognised and described the
“basal grits” of the “Adelaide Series", although
here they have been strongly metamonphised
and intruded by pegmatites. An older base-
ment of highly foliated mica schists, pene-
irated with pegmatites, was observed to
underlie the basal grits in the goree of the
South Para River, and to be separated from
them by a Brass covered, but nevertheless
“abrupt and strongly defined” junction.
Howchin (1926) presented the resulis of
more extensive obsetvationy in this area, in-
cluding a [3 km section extending ENE from
the “Houghtonian” schists in the South Pita
gorec oear the Warren Reservoir Weir,
ihrough the basal grits inte a sequence of
rocks, comprising the Barossa Ranges, which
he eqvated with the lower portion of the
“Adelaide Series". Although clearly recog-
nising the existence of an older basement in
this area, Howchin included within his tasye-
ment some coarsely crystalline schists, such as
Brown & Woodward's highly metamorphosed
Tucks of the Gumeracha Goldficlds, and the
host jocks of the rutile deposits near the
reservoir, which are now known to oaverbie the
“basal grits’.
Hossfeld (1935) refuted many of Howchin's
Views and, although accepting the “basal
grits” south of Williamstown, regarded the
rocks gist al Williamstown os an older
sequence {Woolnough’s Barossian) which
underlay the grits. Miles. (1950) published a
map incerporating the north-western part of
the arca under present consideration and
showed that highly metumearphosed Adelaide
System rocks overlay the haematitic schistose
sandstones, and were faulted against low grnde
Adelaide System rocks belonging to a much
higher stratigraphic level.
Alderman (1942) regarded the high-grade
aluminous schisis and gneisses, and their con-
tained sillimanite, kyanite and clay depostis,
north of the reservoir as products of an exten-
sive metasomatic alteration of original country
kK. 3, MILLS
rocks of regional biotite grade This concep!
of extensive metasomatic activity in the areca
was further adyanced by Campana (1953) and
Campana ef «wl. (1953) in the interpretation oF
the Gawler Geological Sheet. Schists which
Howchin saw as an underlying basement to
the basal grits were interpreted by Campana
as a metasomatic facies of the busal beds.
Hence these rocks were shown as the “Alu-
minous metasomatic zone of South Warren
Reservoir” on the Gawler Geological Sheet.
Campana accepted Howchin's view that the
sequence cas! of this zone could be correlated
with rocks ef the Adeluide System.
Previously (Mills 1963), | authited the pet-
tology of the yarious rock units in the area
south of the Warren Reservoir and considered
that the schists were the products of normal
amphibolite facies metamorphism of rocks of
appropriate composition, doubting the need
for extensive metasomatic activity. The schists
and gneisses south-west of the reservoir were
Observed to underlie Howchin’s “basal grits”,
but at that stage a convincing unconformity
had not been identified. "Whe schistose meta.
sandstone unit became more micactous
towards its base and was thought to pass
conformably downwards info micueeuus
schists and gnetsses.
Whilst studying the mesoscope geometry ol
structures in these high-grade schisis in the
summer of 1967, L discovered that schisis
below the metasandstone unit were much
more complexly deformed than those above,
despite similwritics in lithological character,
and this Jed tu the distuvery of several out-
crops displaying a distinct stratigraphic uncon-
formity between the metasandstone and the
underlying schists, in a section which has
beeome the southern portion of the Warren
National Park, Through personal conmmuni-
cation with Messrs. Ofller and Fleming. this
tediscovered bascment inlier appeared in their
published synthesis of the structural and meta-
morphic. history of the Mount Lofty Ranges
(OMer & Fleming 1968, pp, 248, 252), Since
that time I have engaged in several summer
fleld excursions to the wrea, mappiny in further
detail, and the following account presents the
results.
The area has been mupped using werial
photographs and a base map constructed from
an uncontrolled mozaic, Theré may, therefore,
be slight dislorlions from true orthographic
projection in parts of the mip, bur it has
been found that the grid used on the Adelaide
STRUCTURAL GEOLOGY OF WARREN NATIONAL PARK
und Gawler 1:63,360 Military Sheets may be
satisfactorily imposed. Ft is importint 1
realise Thal the maps, as reproduced here.
present interpreted boundaries and fuults.
Reasons governing these interpretations will
be outlined below, The reader may be dis-
appointed to discover, particularly in the
south-eastern poriton of the areca, that lines
on the map may seem ta have no readily
apparent meaning in the fieki. They are, how-
ever, based on the close inspection and inter-
pretation and careful plotting of al) available
field exposures over the whole area covered
by the map.
Stratigraphic and Metamorphic Retationships
Between Mapped Lithological Units
PRELIMINARY COMMEN1S
The presence of an older Precambrian base-
ment inlier, here referred to as the Warren
Thlier, has now been flimly established an the
hasis of observed wnconformuble relationships
and on structural grounds. Overlying this base-
ment is a mantle of Adejaidean sediments,
beginning with a thick schistose metasandstone
of Aldgate Sandstone type. and passing
upwards into a varied sequence of pelitic,
arenaccous and calcareous beds. The Ade-
laidean =omuantle has suffered three strong
compressive deformations and course recrys-
tallisution in amphibolite Facies metamorphism.
Strong thrusting from the east accompanied
or followed the second deformation and
resulted in several strike Faults with extensive
aggregate movement. The largest of these
faulis hus brought a kyanile-andalusite [acies
sequence of siniple structural chyracter into
conjunction with a cemplexly deformed silli-
manife-muscovile facies sequence, The thrust
slices were folded in a third deformation,
which also updomed the hasement inlier. Fol-
lowing this deformation, the whole area was
uplifted by a very large displacement on the
Williamstown-Mesdows Fault, bringing amphi-
belite facies tocks into conjunction with a
biotite grade sequence, marking the western
boundary of the area. Figure | shows the
subdivision and distribution of lithologicul
units,
THE BASEMENT (Warren INvter)
The pre-Adelaidean basement schists and
gneisses are exposed over an area of approxi-
mate 6.5 km*, These rocks clearly underlie
the cross-hedded schistose metasandstone of
the Adelaidean sequence and are separated
283
[rom that metusandstone unit by an angular
ungonformity, It is proposed that the mame
Warten Inlier be apphed to this basement ex-
posure. The South Para River, downstream
from the Warren Reservoir, has cul a rugged
gorge through this inlier, and the spillway of
the reservoir, like thal of the South Paria
Reservoir further downstream, has been con-
structed on the older Precambrian basement,
There are also Jarge exposures of the basemetit
racks north and south of the South Para
Borge, and in particular at the southern end
of the Warren National Park.
The fresh anificial exposures on the over-
flow race at the southern end of the Warren
Weir provide a conveniently accessible loci-
tion for examining the general nature and
structugal complexity of these rocks. The pre-
dominant rock here is a coarse-grained schis-
lose quartz-felspat-mica gneiss with prominent
lenticular quartzofelspathic und micaceous
compositional layers developed on all visible
scales and displaying various mtnecate fold
structures. A typical specimen is composed of
60% plagioclase (An 14) as unzoned rarely
twinned 0,5-1 mm granoblasts; 10% quartz;
20% biotite as plates up to 3 mm in diameter,
interleaved with some muscovite and altering
to. chlorite; 10% fibrolitic sillimanite, which
has undergone almost complete: sericitisation
and recrystallisation to new decussate muscn-
vite flakes, distributed in lenticular layers; and
the minor accessories rutile, zircon and apatite.
The coarse crystals in this rock are observes
to be mimetic after crenulations in the gneis-
sosity, although there are also pronounced
late strain features, such as bent mica flukes
and deformation lamellae and strain bands tn
ajuartz. The size of the quartz inclusuins its
the plagioclase grains and of accessory zircons
indicate u likely pelitic parentage, There are
rare layers of very coarse biotite rich schist,
and one of these grades into a biotite antphir
holite with accessory rutile, This rock is
composed of about 80% of poorly oriented
interlocking hornblende poikiloblasts 1-2 mm
in diameter, containing about 5% of small
quartz, biotite, epidote, carbonate and opaque
inclusions. The amphibole is pleochroic from
pale lemon to blue-green and is characterised
by strong ratilé exsolution, Thick bioiite flakes
(5%) altering to chlorite are scattered through-
out. The leucocratic fraction fs composed of
equal amounts of quartz and plagioclase (An
45-50) eranoblasts. Rutile, apatite, epidore and
opaque prains are accessories. The textures
13
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STRUCTURAL GEOLOGY OF WARREN NATIONAL PARK
LEGEND FOR
LITHOLOGICAL MAP
im =| Alluvium
( Brotite Zone Rocns.
| Fine-qramed phyllites. dolomites,
dolamihc phyllites, quartzites
HvaNrte-ANDaLusITe ZONE Rocks
Calc-silicate rocks (diopside, scapalita,
amphibole), fine grained khrotted schist
Fine-qrained Knotted schist, pyritie
schist, quartzose schis!
Pink felspathic quartzite horizon
Tramotite rack with interbedded
felspathic and actinolitic sandstone
]
f acy Felspathic sandstone
Siitimanite-Muscovite Zone Rocks
=
tg
S.
si
ey
O
=
Granite qnaiss (quartz-microcline-
Plagicclase-bictite-muscovite gneisal
Course-grained crumpled schist
(kyanite; slaurolite, sillimanite)
Tremolita rack horizon
Felspathic sandstone with Fitanifercus
haematite and muscavite
” BASEMENT
Coarsé-qrained crumpled schist and
Bad gneiss \siflimanife-muscovite zone}
Tertiary deposils and minor intrusions nat -shown
ee Inferred Fault
re
—— Inferred Thrust
en Fault Mélange
1 Le Lenin ty
’ Se
Felice J — Road or Tack
~ River or Watercourse
* Mine or Prospect
Gid Adapted From 1000yd. Military Grid System
Fig. lL. (Opposite). Lithological map showing the
distribution of the principul divisions of
rock types in the region around rhe Warren
Reservoir.
285
and the composition of this rock suggest an
orginal marl. Pods and lenses of quartz and
segregation pegmatites are also common in
the outcrop.
No attempt has been made 16 map litho-
logical variations across the remainder of the
inlier. although exposure is sulliciently good
for this ta be a promising line of attack.
Reconnaissance observations have not revealed
a great deal of lithological variation. Almost
all rocks appear to have had a pelitic or
psammiopelitic ancestry, although there are
some more quartzofelspathic varieties in the
South Para downstream from the weir. The
biotite-rich schists and amphibolites noted a1
the weir are to be found elsewhere as small!
isolated. occurrences, but are apparently quite
sparse. No rocks of the “Houghton diorite”
type (Benson 1909) have been found in this
inlier. The only rocks with an intrusive charac-
ter are scattered quartz pods and dykes and
abundant pegmatoidul rocks, mostly ot a
segregation type but with rarer crosscutting
varieties containing sparse beryl and tourmu-
line.
Amongst the pelitic and psammopelitic
meisses compositional variations range from
sillimanite (sericite)-rich gneisses (e.g. grid
rets. 909082 and 916106), through varieties
with less sillimanite, similar to the weir
gneisses described above, to quartzofelspathic
encisses lacking sillimanite. Some of the latter
may contain abundant well-twinned microcline,
Or puss mto rocks consisting almost entirely of
muscovite und quartz, such as the coarse-
grained crenulated schist immediately under-
lying the melasandsione on the shore of the
reservoir north of the weir. Within each com-
positional group are a number of structural
variants, such as folded, crenulated, lineated,
banded, striped or granulose gneisses, depend-
ing oo the Jocal structural history of each
rock.
A study of the metamorphic minerals and
textures of the basement rocks has indicated
that they were brought to a metamorphic
temperature peak Jate in their structural his-
tory. This metamorphic peak was apparently
uniform over the whole inler and reached
the stage where sillimanite was stable in the
presence of muscovite in aluminous. schists,
No evidence has been found for any extensive
breakdown of muscovite and potash felspar
(mictocline) is confined to the non-aluminous
varieties. Of other index minerals, garnet and
kyanite have been rarely recognised and anda-
236
lutsite aiid staurolite have not so far been
identified, It seems that (he basement rocks
have an apparently similar metamorphic his-
tory to the Adelaidean schists and vneisses
Immediately overlying the metasandstone unit,
and it is helieved that the metamorphic peak
indicated in the basement rocks was identical
to that which may be deduced from tocks of
the Adeluidean mantle. It seems clear that the
pre-Adelaidean metamorphism of the base-
ment rocks involved lower grade conditions
thai those reached in the early Palaeozoic
urageny which affected both basement and
mantle.
Spry (1951) reported a fall in grade from
south to north, involving sillimanite, garnet
and biotic zones, for the early pre-Ade¢laidean
metmorphism of the nearby Houghton Inlier,
which could be construed to fit the above
conelusion, bul Talbot (1962)! has disputed
this grade variation. Talbot (1963) has shown
Ihat the Houghton Inlier has suffered three
metamorphisins. ‘I'wo pre-Adelaidean episodes,
the earlier involving upper amphibolite facies
conditions, and the tater involving pervasive
greenschist facies retrogression accompanying
strong phyllonitisation, were averprinied by a
ereenschist facies (biotite zone! episode accom-
panying the Palacozoie metamorphism of the
Adelaideun mantle. A tentative Rb-Sr date of
867-4492 my, has been suggested for the
upper amphibolite facies. metamorphism
(Couper & Compston 1971), while the Palaeo-
Zoic metamorphism has heen daled al 490415
m.y. (White, Compston & Kiceman 1967).
Jn the Warren Inlier the Palaeozoic meta-
morphism has obscured all evidence of pre-
Audelaidean metamorphic grade in the base-
ment cocks. [| is conceivable that rocks of
the Warren Inlicr have passed through « phyl-
fonilic stage similar to the Houghton Inlier,
bul there & so far no cvidence of at or of any
earlier hivher grade metamorphic episodes,
Che late metamorphic hiktery of the
Warren Inljicr tocks and the overlving Ade
laidewn schists is also similar, involving per-
yasive retrogression of sillimanite to sericite
and muscovite and uf biotite to chlorite, and
she appearance of late strain features such as
hent and kinked micas, lamellar twinning in
plagieclase and deformation bands and lamel-
la@ an quartz.
K. J. MILLS
THE LINCONFORMITY
The actual surface of unconformily is only
clewly exposed in two loeulities; on private
Jand south of the reservoir (grid ref, 925108),
and in the southern part of the Warren
Nuwtional Park (grid refs, 909088-91 2090). The
Warren National Park exposures ate the must
instructive and will be described first.
Several large exposures in the southern
portion of the Warren National Park display
a fingernail sharp unconformity surface, There
is a large angular discordance between the
gently south dipping quartzofelspathic banding
in the basement gneisses and the near vertical
hedding of the overlying metasatidstone
(Fig. 3f). The basement gneisses; here ure
coarse-grained, mica-tich and schistose. The
preminent quartzofelspathi¢ bands may be
related to original bedding, but are probably
transposed and Iack stratigraphic significance.
The basal units of the overlying metasandsione
ure nich in muscovite and strongly schisiose.
These unils become quite friable on weather-
ing, resulting in a ntgative relief against the
more massive basement gneisses, A basal con-
vlomerate is develuped locally at grid ret.
911088. The pebbles range up ro 30 cor in
length, aré moderately angular and are locally
derived from the quartzofelspathic bands of
the underlying basement. The bed is up to
half a nietre thick and i observed to he
welded onto the basement. In adjacent out-
crops the conglomerate is absent and # musco-
vite-vich schistose metasandstonc. characterised
by an unusual abundance of haematite ind ou
few scattered rounded quartz of quartzite
pebbies. to a few centimetres in diameter, is
welded directly on to the basement with a
sharply defined contact. A strong metamorphic
and situctural convergence of the basement
and the overlying Adelaidcan beds has ob-
scured ihis camlact in some exposures. Where
the contact is clear the unconformity surface
is seen to he quite irregular in detail, with
harder bands in the basement protruding into
the basal beds. A metre or so above the
unconformity the bedding in the metasaned-
alone is unallected by these irregularitics.
Following the inlier boundary to the sauth-
west from the above localities, no actual
exposiires of the unconformity were seen,
although a boundary between the basement
! Talbot SLL. (1962)—A study of the structural and metamorphic relationships between older and
younger Precambrian rocks in the Mr. Lofty Rauge Olary Are; South Australia, Univ, Adelaide,
Ph.D, Thesis.
STRUCTURAL GEOLOGY OF WARREN NATIONAL PARK
encisses and the metasandslone can be
mapped, the basal metasanudstone exposures
being agam enriched jn hacmatilc, In the
south-west corner Of the inher the boundary
begins to turn castwards aod disappears
beneath alluvium. Apart from one smul! ex-
posure of cross-bedded hucmatite-rich micta-
sandstone overlying the basement a little
further castwards, no other exposures of the
contact are seen, the southern ind eastern
sides of the inlicr being delincuted by a major
thrust fault.
Following the inlier boundary to the north-
enst from the Warren National Park expo-
Sures, the basal metasandstane beds are
haematite and muscovite-rich, but no actual
unconformity surface was observed. ‘lhe actual
contact lies near the base of a sleep escarp-
ment of husement gneisses. This escarpment
is related to the friability of the basal beds of
the Adeluidean sequence and the erosional
Teststance of the basement gneisses. Rainfall
tun-off trom the basement genciss exposures
engenders a thick undergrowth near the
comlact,
On a track extending porth-west of Lookout
Tower Hill (grid ref. 918105) a haematite-rich
quartz-museayvite rock is seen close to the
inlier contact as Ihe lowest bed in the meta-
sindstone sequence. This rock is composed
of 65% quartz as large grattoblasis to 3 mim,
25% muscovite, 10% haematite, and 5%
barite. as evenly distributed interstitial grains.
Much post-crystalline sttuin ts evident in the
quartz and muscovite.
Following the inlier boundary westwards
around several folds, no actual exposures of
the unconformity surface were found. The
basal beds of the Adelaidean are muscovite-
rich metasandstones, usually bearing notable
hucmalile or a trace of biotite, Bedding is out-
lined by yuartz-rich bunds, und occasional
Tounded quartz or quartzite pebbles to 3 cm
in diameter are scattered along some bedding
planes, The adjacent bastment is mostly a
inieu-rich schist, No unconformity surface was
observed along the long meridjanal western
contact of the inher although the boundary
can be closely mapped. Again muscovite-rich
schistose metasindstones are churactenstic al
ihe basal beds, On the Engineering and Water
Supply access road (grid tef. 902125) a two
metre gap of soil and prass separates whit
appears lo be the basal bed, a coarse hacma-
lite-rich quartz-muscovite metasandstone with
& few small pebbles, from strongly folded
287
coarse-grained quarlz-lelspar-bidtile-muscayile
gneisses of ibe basement. Th haematite-ch
bed can be seen at several points further nor h
hear the inlier contact, but nearer the porthem
tip ef the inter this basal bed has apparently
lensed out and beds of muscovite-rich meta-
sandstone [rom slightly higher in the sequence
come to rest on the basement. From the
northern tip of the inticr (Hale National Park)
to the reservoir the inlier contact was nat
eusily mapped. The unconformity exposure
described by Hossteld (1935, p. 37) could not
be found, although bath busement and meta-
sandstone rocks ure sufficiently well exposed
near the E. & W.S, access road north of the
weir to place the inlier contact on the map
with confidence.
South-east of the weir the fnlier boundary
becomes involved in several tight folds—in
part interpreted previously as ua cross-fauh
(Mills 1963). On the ridge west of Wirrianda
homestead some good exposures of the un-
conformity surface may again be seen. Here
a coarse-grained muscovite and hacmatite-rich
schistose metusandstone overlies a chosely
folded mumuscovite-rich gneiss containing
quarizofelspathic layers and some haematite.
The unconformity surface is again seen to be
irregular in detail, Scattered pebble-like quartz
pods to 10cm in diameter occur in both the
basement and metasandstone and are ap-
parently of segregation ongin. Immediately
south of the homestead the haematite-rich
basal beds of the metasandstone run into the
Wirrlanda Thrust which marks the eastern
boundary of the inlier.
In conclusion, an unconformity between
rocks of the Warrea Inlier and the overlying
Adelaidean sequence is well established, The
inlier boundary can be mapped alung most of
its length with confidence, and details of the
actual unconformity surface can be observed
at two localities. The Jowest beds of ihe Ade-
laidean sequence are composed of schistase
muscovite-bearing nmvetasandstanes and the
basal beds are usually enriched in haematite,
although not as enriched as in ihe ML Bes-
semer sequence on ihe eastern side of the
neatby Houghton Intier (Miles 1950) A basil
conglomerate of the Jocal derivation has been
found wt onky one loewlity.
THE ADELAIDEAN SROURNCE
The siltimanite-muscovite zone
The lowermost units of the Adelaidean
sequence overlying the basement of the
Warren Inlier have reached the grade of meta-
188
morphism where sillimanite and muscovite are
in stable equilibrinm in rocks of appropriate
composition, Vhe basul formation is a thick
brows-hedded = achistose = felspathic metasand-
stone, which is overlain by a thick aluminous
peli Formution, “The top of this pelilic
formation has been cul out by mayor Caults
which have brought the sillimanite-muscovite
zone rocks into conjunction with tower grade
sequences. Three distinet phases of strong
compressive deformation have alfected racks
in the silimanite-muscovite yone and no re-
liable estimates of stratigraphic thicknesses can
be made.
The lithological character of the basal meta-
sandstone formation in Ihis arca has been
carefully described by Howechin (1906, p. 255;
1926, p, 5) whe noted its cluse similarity to
the Aldgate Sandstone, The mectusandstone
supports rather sparse vegetation and usually
forms large lcucoeratic outcrops. ‘Vhe friable
nature of this rock preserves the freestone
tchatacter of the Aldgate Sandstone despiic
high vrade melimurphism, Bedding, cross-
bedding and festoon-bedding, outlined by
laminoe of titaniferous haematite, are beauli+
fully) preserved although considerably ap-
pressed by subsequent tectonic deformation.
The haematite laminae ate normally about one
millimetre in thickness, but rare layers up to
10cm in thickness huve been observed, Miles
(1950) presents the results of a chemical
analysis of the ulaniferous haematite from at
specimen collccted in the northwest corner
of this area, In some oulcrops (eg, grid ref.
908070) the metasandstone is strongly mag-
nec; Magnetite presumably replacing the hae-
mate us the principal opaque accessory, H
is not known whether this magnetic propery
is confined to certain beds.
As described in the previous section, the
lowes beds against the unconformity are nor-
mally enriched in haematite, this component
being scattered evenly throaghout. These hae-
matite-tich beds are not mare thin a Tew
metres in thickness aid in roahy places they
thin ovt and disappear entirely, The next
30-50) tnetres of section consisls of a museo-
Vite-enriched metasandstone, perhaps best
desccibed os a quatlvamuscovite schist, The
remainder of the sectiun, perhaps amounting
ty SOO metres, consists of Selsputhig metasand-
stone, characterised by titaniferous hacmatite
Inminae outlining the bedding surfaces, with
lutenmitrent imuscoviteenriched layers. The
base of this mere qulittzofelspathic section
K. J. MILLS
was taken as the base of the psammutic forma-
tion oo my eurlier map (Mills 1963). As
desevibed by Howchin, rounded pebbles up Lo
30em in diameter, mostly of quartz or fine-
graihed gqualtzite. ace distributed throughout
the sequence, ullhough more commonly eo-
countered at certuin horizons, True conglo-
meratic beds are rarely observed, the pebbles
being mostly scattered unevenly on bedding
surfaces throughout the outerop. In some
exposures the pebbles are sten to have suffered
a strong flatlening and some elongation result-
ing From tectonism. Stratigraphic bedding can
usually be identified and opportunities for
facing observations based on cross-bedding
are numerous. Some slicing and transpasiiin
of bedding has heen woticed in the muscovite-
enriched section near the base of the stqtience
in. the South Para Gorge, Occasional pegmi-
tite and milky quartz veins carrying accessory
ilmenite plates Intrude the metasandstone
throughout the area.
Sixteen specimens covering the composition
range of the psammites were examined micro-
scopically. Quartz, ranging from 25-75%,
occurs in some rocks os Jarge strained clasts
up to 3mm. while other suniples, particularly
the qwore micaceous ones, demonstrate all
slages in the recrystallisution of original
strained clasts to new metamorphic grano
blasts. This recrystallisxtion is closely nsso-
ciated with the axial surface schistosily of the
second compressive deformation. In some
samples the new quartz grains preserve a pro-
nounced preferred orientation of their c-axes,
wppurently related to the second delormatiun.
Sume quartz clasis are sagenilic. Potash felspar
(0-30% } is the sule felspar component. It
usually occurs ax strained tartan-twinned cfists
up to 2 mm. In some specimens the clasts are
apparently untwinned but have partially re-
crystallised to small well-twinned pranoblasts
near their margins, Muscovite (S-50%) is pre-
sent in all specimens, usually as large flakes
vp to 2mm, commonly showing late strain
effects. It appears to be wholly of meta-
morphic origin and is. responsible for the pre-
servation of many of the tecionicully imposed
features of these rocks. Biotite is present in
many simples up to 5% and is usually pleo-
ehraic olive brown to pale lemon. Opaque
grains (1-10%) are invariably present. Apatite,
Zircon, monazite and tourmaline, pleochroic
pale pink to olive green, are accessories. Con-
sidering the coarse-grained micateous Entisses
above and below the schistose sandstones, the
STRUCTURAL GEGLOGY OF WARRTIN NATIONAL PARK
preservation of quartz and felspar clasts at
this metamorphic grade is quite remarkable
and the weakly metamerphosed appearance of
the more quartzofelspathic varieties of this
metasandstone miist be partly attributable to
the preservation of these clastic features,
The upper contact of the metasandstone
ligains! the overiying sillimanite bearing pelitic
schists is shaeply defined. In the south a thin
intermitient tremolite rock bed, aormally ex-
pressed at the surface as an opaline replace-
ment rock, tharks the cantact, while in the
east sume pelitic sehist is inserted between his
bed and the metusandstone, In the north-west
a sliver of coarse-grained sillimanite bearing
pélitic schist is observed to rest directy on the
schistese sandstone, The frue nature of the
tremolitie marker bed can be observed in
debris from a shaft sunk on the ridge nor.h
of Sailors Gully where the rock below the
weathered profile is composed of interlocking
pale-green trenalite-actinolite prisms, Else-
where the surface expression m an opuline
ohaleeduny showing tremolite casts and some
lale.
Vhe thick pelitic formation above this tre-
molite bed has been previously described in
some detail (Mills 1963, p. 171) These rocks
have auiicred intense folding and compression
and true bedding cannot be recognised with
cerluinty.. Although apparently quite thick, a0
estimites oO! straugraphic thickness can be
made. The socks are coarse-grained micucenus
schasts.ind gneisses, usually displaying alumino-
sificnte knots, tross-cutting crenulatiens and
small folds in their schistusity, leading to the
field name “crumpled schists” (c.g. Miles
1950), ‘The major primary constituents are
quarty, plagiaclase (An 0-15), biotite and
muscovile with variable amounts of sillimanite
and kyanile and rarer staurolite and garact.
Assemblages of these imtinerals developed in
rocks of appropriate pelitic composition and
approached equilibrium’ in a sillimanite-
Inuscovite grade metamorphic peak during the
second deformation, As wilh the basement
encisses, HO Metasomatism is considered to be
necessary for their formation. A pervasive
retrogression, which took place in the inier-
Kinematic periad between the second and
third compressive events, has resulted in the
extensive alteration of aluminosilicute minerals
and biotite to sericrte and chlorite. This retro-
gression is more pronounced in some zones
where shearing associated with thrust move-
muuts is belicved to have been important.
289
Some introduction of H.O may have secom-
panied this tetrogression,
fia the eust, part Of the pelitic formation has
been replaced by a hody of granite gneiss, as
pieviously described in greater detail (Mills
1963}. Although the chemical processes which
led to the formation of this granite gneiss are
hot yet understood, extensive chemical migra-
tions are not envisdged in its formation. This
eranite gneiss and its immediate envelope of
schists have escuped retrogressioa, Ti is en-
visaged that the granite gneiss body behaved
as @ sohd impermeable block durmg the reire
sressive cpisode which affected the adjacent
schists,
In Conclusion, the hasal metasandstones and
the woverlying pelitic schists have been sub-
jected lu a metamorphic peak in the middle
amphibolite facies under temperature condi-
lions where sillimanite was in equilibrium with
muscovite. Some clastic textures have been
preserved in the more quartvofelspathic meta-
sandstone beds but in the pelitic schisis exter
sive recrystallisution took place, Sillimanite,
kvanite, Staurolite and garnel were crystallised
during this. metamorphism aod grannisation
occurred Jogally, At a liter sige under yreen-
schist facies conditions. extensive relroyression
took phive in the pelitic schists.
The kyanite-andalusite 2one
A sequence of Adelaidean rocks belonging
to the lower amphibolite facies have been
(aulled against rocks ol the sillimunite-musco-
vite zone in the south, cast and north-west.
North of the Warren Reservoir, Alderman
(1942) placed these rocks in the bjotite zone,
a reasonable supposition considenng the abun-
dance of biotite-quartz-felspar schists which
oceur in this sequence. Aluminuus beds con-
taining undalusile or kyanite Knots are rare
and these koots are usually severely relro-
gressed jo sericite, Cale-silicaie beds in the
seqttence carry diopside, scapolite, amphibole,
epidote and oligoclase-undesine, confirming
their amphibolite facics character. This se-
quence of rocks is characterised in the field
by the fing to medium grain-size of the pelitic
and psammopelitic umnils and their stmple
structural character dominaicd by a single
schistosity, which in most outcrops is parallel
or nearly parallel to well-defined bedding
planes and bedding laminations. Crenulations in
this schistosity are rarely found, and if present
are weakly developed. The sequence ¢ast of
STRUCTURAL __MAP
OF THE
WARREN NATIONAL PARK
AND WESTERN POATION OF THE
MT CRAWFORD STATE FOREST
O00 ine Yet
UKAPRAINGA
Mine |
=
ir
9
=
Zz
e
r
#
Boe
ee
o-_
=
Le
z —
RUST ASS
peek?) fl =
aa i 28
rae I aN
ey r vA s “
—=
STRUCTURAL GEOLOGY OF WARREN NATIONAI. PARK 291
LEGEND FOR
STRUCTURAL MAP
Lithological contacts _
oo a ad
Trends of bedding surfaces
7
Unconformity
ae
Marker ‘horizons
Stratigraphic facing = we
-es tablished
—_—_—~
Foults ~inferred
-inferred
are
Thrusts
-established
aa
Fault mélange ress
ADELAIDE SUPERGROUP
BEDDING eS = *
Sq Inclined Vertical Horizontal Inverted
FIRST DEFORMATION
\ Si, %,
5, LiBlg* Ls,
SECOND DEFORMATION
Sz 4, 52
S, LIBIg? \LIB)e?
THIRD DEFORMATION
% Sa S. Sa
Sy Xk LiBles Bg? L(B)e? \
BASEMENT
Compositional layering »
Schistosity *
Axial surface to folds in schistosity “s
Lineation in schistosity ™
Hinge lines of folds in schistosity
Fig. 2. (Opposite). Structural map showing the
principal structural features. in the region
around the Warren Reservoir.
the granite gneiss has been briefly described
(Mills 1963, p, 171).
Exposure of this sequence is very poor iin
the south and south-cast bul it seems that the
lowest unit is a pink meta-arkose containing
sorne haematite laminac, but no mica, and
preserving good clastic churaclers, South of
Watts Gully this meta-arkose is overlain by
a sequence of fine to medium-grained quartz-
felspar-biotite schists containing some pink
microcline-rich meta-arkose beds with acces-
sory pyrite. Then follows a tremolite rock
marker unit, expressed at the surface asa ten
metre width of opaline replacement rock. This
unit has been traced intermittently, using
exposures. of opaline replacement rock or talc,
around to the eastern side of the granite gneiss
where it forms a thick and useful marker
horizon. At the surface it is usually expressed
as opaline chalcedony, but fresh samples from
below the weathering profile consist of coarse
interlocking tremolite prisms. North of the
present area this same horizon is apparently
represented as a tremolite marble and has been
traced from well north of Williamstown to. the
proximity of the Warren Reservoir (Howchin
}926, pp. 7-8; Hossfeld 1935).
East of the pranite gneiss the tremolitic
marker unit is overlain by a mica:schist group
consisting of fine to medium-grained quartz-
felspar-biotite-muscovite schists with thin inter-
bedded meta-arkoses, Most of these meta-
arkoses are fine-grained, pyritic, graphitic and
very enriched in microcline. A useful pink Sels-
pathic quartzite marker horizon crops out
persistently a little above the tremolitic marker
unit. Some mica schist beds contain small
knots which have altered to fine sericite. Rare
kyanile relics have been found in these knols.
The uppermost part of the sequence examined
is a Calc-silicale group consisling of inter-
bedded fine to coarse cale-silicate rocks and
knotted fine-grained mica schists.
A wedge of rocks belonging to the kyunite-
andalusite zone has been mapped west of the
Ukuparinga Copper Mine, in the north-west
corner of the area. The lowest bed here is a
thick medium-grained dolomitic marble, which
may be equivalent to the tremolitic marker
bed east of the granite gneiss. Specimens of
this rock consist largely of dolomite with
variable amounts of tremolite, talc, muscovite
and primary chlorite. Calcite is rare or ubsent-
‘Two. analyses of this marble are presented by
Miles (1950), This marble is overluin by a
mica schist sequence, containing fine-grained
242 K. F. MILIS
pyritic meta-arkose beds, very similar to that
east of the granite gneiss. Higher in the
sequence some mica schists have sericite knots
containing andalusite relics. The cale-silicate
group has not been recoguised here, although
a second marble unit crops out high ia the
sequence against the Williamsiown-Mecudows
fault.
Biotite zone
No attempt has been made te map beds in
the low grade sequence expasedl west of [he
Williamstown-Meadows fault. Near the [fault
the main rock types are fine-grained grey
dolomites, grey-brown dolomitic phyllites,
phyilitvys and fine-grained quartzofeispathic
pyritic metusiltstones. Rocks of the biotite
zone sre usually extremely fine-grained
(0.01-0.03 mm). A pale mustard-brown weakly
pleachraic biotite is characteristic of the phyl-
litic tocks. Dolomite seems to dominate over
calcite in’ the carbonate rocks. A thick
sequence of grey impure dolomite; (20%
quartz, plagioclase, muscovite and opaque
impurities) is exposed south af the South Para
River
CONCLUSIONS
‘The basement rocks of the Warren Inlier
are Wnvenformably overlain by the Adelardean
rocks. The Adelaidean sequence is divided into
three portions belonging to different mela
morphic and structural levels by several major
fault surfaces, The basal sillimanite-muscovite
zone rocks and the kyanite-andalusite zone
tocks of the Adelaidean sequence are sepu-
rated in this. area by Faults which ate surfaces
of marked metamorphic and structural di-
continuity. It therefore seems likely that there
is a large stratigraphic gap between the
Sequences represented within these meta-
morphic zones, Further detailed mapping, par-
ticularly to the north of the present area, may
help to determine how much of the struti-
graphic sequence has been removed. Until this
problem is solved it seems unwise to attempt
correlations of individual beds or parts of the
sequetice with other beds or sequences in the
Mt. Lofty Ranges. The Williamstown-
Meadows fautt brings biotite zone rocks in
the west against sillimanite-muscovite zoue
rocks. As suggested by Miles (1950) the biotite
zone tocks probably belong to a much higher
stratigraphic and metunwrphic level in the
Adelaidean sequence and displacernent on this
faull must be very large.
Structural Relationships between Mapped
Lithalogical Units
PRELIMINARY COMMENTS
Three successive compressive deformation
events, Fi, Fe and Fs, have affected the Ade-
Juidean sequence in the vicinity of the Warren
Reservoir and the structural features produced
in these de‘ormative phases will now be
described more fully. Metamorphism of the
Adelaidean sequence began betore or during
Fy and reached a peak temperature in the
amphibolite facies early in the Fy episode.
A series of east to west thrust movements
occurred Jale in the F, event and extensive
Telrogression under greenschist facies condi-
fens accompanied or followed these moye-
ments. ‘Uhe thrust sheets were folded during g
Tather hrittle low temperature F, event. After
F, and late in the cooling history the Williams-
town-Meadows fault developed its maximum
displucement,
In describing the struelural features and
events in the Adglaidean scquence it is cane
yenient to introduce the following shorthand
nojatian? (see also map legend, Fig. 2)—-
S) —surface of compositional
layering relating to pri-
maty bedding.
F,, Fu, F, --the three successive
compressive events.
S,- Su. Sp —axial surlaces to fulds
produced in the F,, Fa,
F; events respectively.
—fSulds in bedding with 8.
as axial surface etc.
—lineations resulting from
Sp-Sy intersections etc,
—strain elongation axis in
Sy ete.
Orientation measureinents are recorded in
telation to true north-point azimuth. Figures
2 and § illustrate the principal structural
features
B(Sp-So) ele,
L(Sy-So) ete,
LiS,) ete.
THE BASEMENT (WARREN INLIER)
At the type locality of the basement rocks
at the Warren Reservoir Weir a partial history
of tectonic events may be dcicrmined. As in
all exposures of the basement rocks, bedding
2 Owing 10 ptinting difliculties this form ef notation is adopted im the cext cather than ths more
conventional form shown on the map Jegend.
STRUCTURAL GEOLOGY OF WARREN NATIONAL PARK
with stratigraphic Significance cannor be
proved, although quartzofelspathic bands and
layered lithological Variations may reHect 4
primary bedding precursor. Lithological layers
and schistosities in Ihe expusures near the weir
are strongly folded on all scales and at first
sight the folding appears to be rather regular
wilh a prominent schistosity. defined by
course mica plates, crenulated and folded
about shallow plunging axcs (Mills 1963). This
schistosily is observed in detail to be a highly
evolved crenulation cleavage, and is therefore
an axial surface schistosity belonging io at
least a second deformation event. In paris of
this exposure this schistosity has been folded
into recumbenr Z-shaped folds with near hori-
zontal hinge surfaces, and these folds have
been overprinied in turn by a widespread later
folding with a moderately west-dipping axial
sutfuce, produging ¢renulations in the earlicr
schistosuy with ow wavg¢lengih of O,5-3 cm,
Thus, at least four overprinted fold even's
can be distinguished in this exposiire, and
there are yet other folds present which may
represent a lifth deformation. The earlier faleds
in the weir exposure tend to huve a. ptygmatic
style in which the fold hinge lines have a more
regular orientation than the hinge surlaces,
Nevertheless, the hinge lines remain curvi-
linear on all scales and the folds are never
strictly eylindrical, The Jater folds are also
irregular and often of polyclinal siyic, this
being puriistly induced by the presence of
many large intrusions of pegmatite and blocks
and lenses of quurtzolelspathic gneiss which
have acted as. buffers inducing inhomogeneous
strain during the later fold movements (Fig,
3b, c, e). Due to these irregularities, reliable
field? measurements of the orientulion of siruc-
tural elements are difficult to make, even in
the most favorable exposures.
Lisewhere in the inlier most exposures show
signs of complex folding and overprinted de-
formation structures, justifying field names such
as crenulated or crumpled schists. Bedding
cannol be veriligd and any lithological layering
present is strongly lenticular und suggestive of
Irunsposition processes, Sone exposures are
very irregularly deformed and tio orientation
measurements can be made, but some of the
more micaceous schists display fairly regular
folding of their schistosity, approaching local
cylindricity, There ts much variation in the
style of small folds from concentric to shar}
chevron folded forms (Fig. 3a. dc. fi). Most
pegmatite intrusions have alse been folded
243
along with the host rock schistosnies.
Examination of exposures in the vicinity of
Lookout Tower Hill agam suggests multiple
deformation involving at tcast four conipres-
sive eVents, the Jutest phase involving ubi«
quitous tight to ope chevron style folds in
the earlier schistosily of the more micuccous
rocks.
Microscopically the basement socks are
cuoumegrained, wilh a prominent schistosity
defined by oriented mica flakes, which are
oflen segregated into Jensoidal gneissic layers
with signa of evolution thtaugh an earlier
crenulation cleavage stage. In parts of the
inlier a strong mineral fineafion, detined by
sillimanite fibres in aluminous schists or mic
plate dimensions, 18 observed within the <chis-
tosity, Folds and ercnulations in the schis-
(osily are ubiquilous and are mostly pre- or
synmutamorphie. Some ot these lolds seem to
be closely related to the F.,, deformation in the
overlying metasandstones. Weak Tate meva-
morphic strain features, undulose extinction
und deformation lamellae in quartz and kink-
ing and bending of mica plates, are observed
In Most specimens,
Although some structural measurements
have been made in the basement rocks (Fig. 6,
Sub-areu Aj, these have little or no meaning
in terms of the complex structural history 14
which these rucks hive been subjected, and 4
more complete understanding of their -struc-
tural history may. only evolve through a more
detailed structural analysis of the whole inlier,
‘THE ADELAIDEAN SR@URNCE
Yhe sillimanite-muscovite zane
The busal unit of the Adelaidean sequences
is & schistose metasandstone in whith quartzo-
felspathic and mica-rich layers and titaniferous
haemittite Jaminae clearly oniline prinnury
bedding (So), and well preserved cross-beddiing
structures provide evidence for strahyfaphic
facing as described previously. Plates of white
mica, presumubly crystallised from a former
clay/felspar fraction in the sandstone, ate
present in almost all beds of this unil but
become more abundant towards the base.
‘These platy crystals have enabled various tce-
tonic structures to be preserved m the aeta-
sandstone, and [rom a study of the mica plate
orientations a lectunic histury ¢an be erected,
Like the overlying pelitic scquence the basal
meétasandstones have been ifected by three
major deformative events F,, Fy, and Fy
(Pig, 6). The F; ond F, events caused tight
794 K. J. MILLS
folding of the mctasandstone throughout most
of the area considered here. The Fo event in-
volved sirong compressive deformation near
the peak temperature of metamorphism and
this combination has tended to obscure the
effects of the F; event im most exposures.
During the Fy event a strong S$. schistosity
evolved through the crehulation and trans-
position of an earlier S, schistosity. The effects
of the Fy event are rather weak and are
responsible for some post-crystalline crenu-
lations in, earticr schistosities in the centre of
the area and macroscopic folding of earlier
structures in the south-east (Fig, 6, sub-arcas
M, N, 0, P, Q).
Most cxposures of the metasandstone: dis-
Play a schistosity defined by oriented mica
plates. In the less micaceous rocks same
difficulty is experienced in differentiating this
schistosity as S, or Su. In some of the more
micaceaus tocks S,; and S, are both present
and can be clearly distinguished on the basis
of overprinting relationships, Where S, is most
clearly preserved it is seen to be defined by
well-oriented mica plates. The mica crystals
are grouped into anastomosing clusters of
nearly parallel plates separated by a single
Jayer of quartz and telspar grains. The clusters
are evenly distributed rather than forming the
segregated trains of mica plates more charac-
STRUCTURAL GEOLOGY OF WARREN NATIONAL PARK
lertstic of The Sy microstructire, The gram size
of the mica plates does not seem lo be a
consisicnt leatyre for the distinctian ef §,
ant S.. In situations Where (te S, surface lies
at a distinct angie to Sp and in cross-bedded
micaceous melusiodstunes, S, can be shown
to be an imposed tectonic feature and not a
“bedding foliation”, However, in most out-
crops 5, is sensibly paraliel to S,, suggesting
a position on the limbs of very appressed F,
Falds,
Owing to the strong overprinting effects of
the Fs event, inacroscopic F, folds cannot be
readily mapped oul, but the shape of the base-
meal exposure sugvests the position of some
macroscopic F, folds, A stereographic plot of
poles to bedding surfaces measured over the
large synclinal structure straddling the cenire
of the inher results in a broad girdle pattern,
the pole of which, plunging ut 20° towards
202°, may be laken to represent the mean
macrascopic B(S,-S,) fold axis for this strne-
ture (Fig, 4, sub-area L), The orthographic
block diagram of Fig. 8, showing the general
form of the stnicture, has been derived using
this calculated H, uxis and the local orienta-
tion data on the overprinted F, and Fy struc-
tures. In the north-west of the area, for
example along the Engineering and Water
Supply track crossing lhe metasandstone on
the western side of the inlier, same mesoscopiz
F, folds show tight appression wiih ihe 8,
axial surface structure striking 002° and dip-
ping steeply eastwards. Insufficient observa-
298
lions have been made bo discuss strain in the
PF, deformation,
AML stages in (he Gevelopment of the S.
Stvucture can be observed in the metasand-
stone, from faml, open crenulations in the
earlier S, schistosily, to uppressed crenulativns
in which the mica plates have become con-
centrated in the limb regions, leaving. (he
hinge areas enriched in quartz and felspar with
only a few diversely oriented mica plates; to
& new gneissose schistosity in Which the mica
plaies are largely segregated into trains
parallel to So, leaving little sigu of the previous
S, structure in the rock. In the more quartzo-
felspathic beds both the quartz and lelspar
grains have discoidal shapes with Jong axes
lying in the Sg surface, The mica plates defin-
ing the crenulations of the new S. schistosity
characteristically lack pronounced — strain
features and evidently underwent recrystallisa-
tion during and after the Fy, event, that is
syntectonic and podst-tectonic crystallisation,
and ihis feature serves to distinguish F,, crenu-
lations from Fy crenulations over most of the
area, However, in the south-western portion
of the Warrén National Park detormation in
the Fa event outlasted recrystallisation. Here
mica plates huve been strongly kinked and
intragranular movements outlasted strain
relieving processes. Relict quartz clasis shuw
strong wndulose extincnion with deformation
bands parallel to S, and newly crystallised
quartz grains have developed a pronounced
caxis fabric during the F. event. This fabric
Fig. 3, Structures in basement rocks of the Warten Inlier and at the base of the Adelaidean succession,
a—Prulile of folds in coarse-grained quartz-mica gneiss, Warren tnlicr, A well developed
gneissic schistosilty has been thrown into regwiar chevron folds ina deformation: which may be
correlated with Fs in the overlying Adelaidenn succession, Traced from photograph. Location
918098.
h,—Profile of folds in basement rocks, Warren Reservoir weir. Quartzofelspathic layer in mice
unecixs stuwing polyctinul folds. A crenufation cleayage cross-cuts the folds (shading) aad
crennlates an earlier axial plane schistosity in the mica gneiss. Freehand sketch, Location
916116,
c and ¢—Disharmonic folding of quarizofelspathic layers in coarse-grained mica gneiss, Warrett
Reservoir weir, Small folds associated with the crenulation of an earlier schistosity show 2
polyclinal style. Traced from photographs. Location 916116,
d.—Profile of open folds in course-grained mica gnedss containing thin quartzofelupathic luyers,
Warren Inlicr, ‘raced from photomicragraph.. Lacation 928098, ,
f. Sketch of unconformily between basament rocks of Warren Inlier and basal motayandstone
of Adeglaidean succession. Drawn looking eusiwards. The quarizufelspathic basement (stippled)
has thin ovitaceous layers dipping al a low angle to the south. The gaconformity is steeply dipping
but has an irregular detailed topography, A hacmatite-rich mic¢uceous. metasandstone showing
traces of bedding in the form of hacmatite-rich seams (fine stipple) is welded onto. the base-
ment und has a strong S, schistosity, delined by the orientation of mica plates, dipping to the
south (shading), Freehand field sketch, Location 912091, Warren National Park,
“—An angular quarts pebble immersed in coarse-grained Muscovite-rich melasaadstone near
the hase of he Adelaidewn succession. Note the Fy crenulations im the earlier S$, schistosity-
Vraced from photomicrograph. Location 908107.
f.—aAn example of the uverprinting of carly folds in schistosity of a quariz-muscovite gneiss
by later folds, Warren Tnlier. Sketched from hand specimen, Location 917106,
296
has not yet been investigated in detail, but
appears. 16 involve a girdle or crossed girdle
pittern in which the cakes tend to be sym-
metrically disposed to the Sy surface and iu
the strony L(S,-S.) lincanon lying within jt,
A lineation, L(S,-8.), is scen to lic in the
S. surlace in most exposures throughout the
area. Where 3, is developed only as a crenu-
lation cleaviize this lingation is the crenulation
axis and is the geometrical result of the inter-
secion of the §, and So surfaces. Where
Sx is more strongly developed into a new
schistosity the lineation may take the form of
a mineral streaking effect on the S. surface,
Microscopically this. mineral streaking is seen
to be the visible effect of elongate mica putes,
with variable dimensions but averaging 1:5710,
whoch are inutuzonally arranged about the
lineation axis with their longest dimensions
parallel to the lineation, Some quartz and fel-
spar grains are also elongate parallel to ihe
lineation. Quartzose pebbles are often discard
within S, and slightly elongate parallel to the
lineation, However, whether this Jineation,
which hax heen mapped as an S,-S, inter-
section lineation in the field, is alsa a principal
axis of strain has not yet been determined.
Most of the mapped macroscopic folds
within the metasandstone and involving the
unconformable contact between the metasand-
stone and the basement rocks have proved lo
he Fy folds. Mesoscopic F, folds in both
bedding and S, surfaces are not uncommon.
Two small B(S,-S.) folds from nomh of the
well, illustvated in Figs. 4a and 4b, were
analysed using the technique sugycsted by
Ramsay (1962; 1967, p, 413) for unravelling
flattened flexural slip folds. In both causes the
western lintbs proved to be more “‘flatiened"
than the eastern limbs, und a mure micaceous
layer had both limbs “flattened more than
100%", These resulls are unrealistic and
indicate that olher mechanisms besides flexural
slip and flattening were involved. The grealer
“flattening” of the western limbs could he
explained by a component of simple slip or
sheacting motion, involving intergranular
movemeits, which would be consisient with
the enst to west thrusting movement charac-
teristic of the laler stages of the F. event.
In the incompetent micaceous layer mass
movement of material from limbs 10 nose
could explain the anomalous “fiattening" in
these layers. However, 4 more generalized
mechanism may have heen responsible for
these folds (Hobbs. 1971),
K. F. MILLS
In the vicinity of 906093, crenulations with
a Wavelength of 4-1 cm are seen to huve hinge
surfaces parallel to those of farger folds.
These hinge surfaces suike at 335” and dip
steep westerly and are interpreted as belong
ing to F, folds. In some specimens L{S,-S.)
lineations ace seen to be folded over these
crenulations, Thin sections show that all
crystals have preserved intense strain features
related Jo this deformation, In the south-vast,
in the vicinity of the Gumeracha Goldfielkts,
struclures in the metasandstone have been
folded aver a large Suuth-easterly plunging F,
anliform,
In thin section, specimens of the metasand-
stone from throughout the area show some
strain in the quartz and mica crystals, but it
is not known whether this is reluted to the Fa
deformation event outlasting reerystallisution,
the F, event, or some other luter mild phase
of deformation.
Rocks of the thick pelitic sequence over-
lying ihe basal metasandstones are charac-
terised by a coarse gtain size, a pronounced
schistosity defined by mica plates, and an
abundance of mesoscopic linealions and lold
structures. Like the basement gneisses, there
is much variation in the textural and sirucg-
tural features of each rock depending on the
original composition and the Jocal tectonic
history. Figures 6 and 7 summarize the meso-
scopic orientation data measured in the Ade-
laidean sequence, Similarities in the orientation
data measured in the metasandstones and
overlying schists und gneisses may be noted
(e.g. cumpare sub-areas R ind H with sub-
area J).
The most attractive structures within the
pelitic schists are the open tu moderately tight
mesoscopic folds belonging to the F, event,
These folds show varying degrees of develop-
ment in different oulcrops and are apparently
ahsent in some parts. The style of the F,, folds
and their patchy distribution suggests that they
were produced in a rather mild Jale lestonic
event when compured with the intensely bonm-
pressed nature of the carier siructires. The
main schistosily in these rocks can be observed
to be an axial surface slructure to very ap-
pressed, almost isoclinal, folds and to be a
highly evolved crenulation cleavage belonging
fo the F., cvent. No mesoscopic F, folds have
been identified in these schists, although relics
of an earlier schistosily testify to the presence
of a possible axial surface structute, equrva-
STRUCTURAL GEOLOGY OF WARREN NATIONAL PAKK
lent to slaty cleavage, for this earliest tectonic
event,
No bedding with stratigraphic significance
has been proved within these schists, Len-
ticular’ compositional layering, labelled $,. is
observed within the larger exposures, but even
this thickest layers are not traceable as bedded
units, Attempts to map out lithological unite
ona larger scale, using compositional and tex-
tural variations, have failed. This could be
partly cue fo luck of silicient continuous
exposure, Relics of an carly schistasity, S;,
presumed tu be wn adial surface structure for
the F, event, are to be found in many expo-
sures, These relics take the form of an early
schistasity acting as the folded surface in
some woclinal Fy folds, which are usually best
preserved in rocks of more quuttzase cam-
posilion, and relics of an early crenulated
schistosity in microlithons within the F.
microstructure in rocks of more micaceous
composition. The observalion of disc-shuped
quartz-sillimanite and staurolite knots lying
within the S, surface forming the folded sur-
face of F, folds suggests thar these minerals
erew during or after the F, event and before
the imposition of the Fe event. No B(Si-S1)
foids have been identidied in the schists. From
the secant evidence available it seems that 5,
and S, were nearly parallel in these rocks,
and this could be attribulable to a sitwation
on the limb of a large appressed F, fold.
The mai schistosity in these rocks, Ss,
lonms the axial surface structure to tight folds
produced in the Fy, event. In some exposures
relics Of appressed crenulations in S,; with
sharply attenuated hinges are observed within
the §, microstructure. but transposition of
the earlier schistosity has been so great that
only rarely are these crenulation hinges. pre-
served, The more fully evolved S. micro-
structure is characterised by w coarse-grained
gntissic texture in which quartz and ftelspar
granoblasts are sezregaled into thin lenticular
plates interleaved with trains of coarse-grained
mica Makes. Mies plutes comprising the mica
(rains commonly show a consistent imbricate
Or echelon stuckiny arrangement indicative of
an evolution of the Sy microstructure through
the concentration and metamorphic segrega-
fon Of a pre-existing $, mica preferred
orientation (Fig. 4g). Isolated mica plates
withic the quurtzofelspathi¢ Jenses tend to
show a diversity of orientation, Where un-
affecied by the Fy event the So surfaces in
297
tocks south of the reservoir have a mean
strike of 350° and dip 75° east.
Fulds in compositional layering and 5,,
with S. as axial surface, are not vncommenty
met with, bul are less obvidws than the later
steep plunging F, folds. These F. folds are
tightly appressed and almost isoclinal with
thickened hinge regions (Figs. 4c, d, c, fy
They are overturned to the west and ther
hinge lines plunge at variable angles. within
Sq. The strong S, axial surface structure tends
to obliterate the folded S, surfaces, especially
in. rocks containing coarse porphyroblastic
knots, A strong lineation L(S,-S,) results from
the intcesection of the 5S, and Sy surfaces, This
lineation takes the form ol a mneral streaking
or rtodding defined by ellipsoidal shaped
granoblasts and mica plates, or is more rarely
Aven a6 crenulation hinge lines. The lineation
is usually parallel to the axes of Fa lolds itn
S) or S,, bul examples are known where the
lineation is more steeply plunging than
B(S,-S2) fold axes, suggesting that S, and S,
were not sttictly parallel before the Fy,
deformation vent. Near the old reservoir
rouwd bridge the L(S8,-S,) lineation has a
shallow north or steep casierly plunge, bul
further south und west a moderate south-
easterly plunge becomes predominant, Cross
joints have been developed perpendicular to
this Imeation. It is not known to what extent
this lineation is also a principal axis of strain.
A mild Fy event occurred lute in the meta-
morphic history producing patchily developed
open to tight F, folds, These folds are mostly
B(S.-S,) folds, but locally B(S,-S,) and
B(S,-S,) folds may be found. Several
examples of the overprinting of Fy folds on
F, folds have been observed, The F; folds
usually have open concentric, boxlike or
V-shaped styles, which pass into regular or
polyclinal chevron falds in the more strongly
deformed exposures | Fiz. 5). Mesoscopic folds
near the reservoir haye wavelengihs up to
6 metres, but much larger macroscopic folds,
auch as the Gumeracha Goldfields antiform,
occur in the south. A mapped macroscopic.
told in the south-west has a curious irregular
form in which S, has been folded about ah
axis plunging 20° to 153°, but the F, struc-
tures show various. anomalous -orientutions.
B(S.-S;) folds plunge steeply north-cast to
south-east near the reservoir (Fig, 7) und
moderately south-east in the Warts Gully
region (Fig. 6). Rotation of earlier L{S\-S,)
lineations in conical surfaces Suggests 4
288 K. J, MILLS
Fic. 4-
STRUCTURAL GEQLOUY OF WARREN NATIONAL PARK
fexural slip mechanism. The Fz folds tend to
split alang their hinge surluccs on exposure
and this is ofterr the only expression of an
Ss wxial surface structure. However, in the
tightest F, folds an S$, crenulatwon cleavage
has begun 16 appear, Fy folds tend to have a
polyclinal style with considerable variation in
the orientation of their axial surtaces, Neal
the reservoir the Fy folds have a left-handed
vergence and §., axial surfaces vary tn strike
from NE-SW for open fold styles. to N-S for
appressed styles where the S; crenulation
cleavage has begun ta appear (Fig. 7). In the
Wults Gully tegion the folds have a right-
handed vergence, the S, axial surfaces in the
open folds having a NW-SE strike swinging
1o N-S as the folds become tighter (Fig. 6,
sub-sreas M, N, O, P, Q). Under the micro-
scope lhe F, folds near the reservoir seem
to show a greater degree of healing of both
quartz, and mics in their hinge tTegions than
do Fy folds in the west and south-west of
the area, ‘Thi may suggest a slightly higher
lemperatute during the Fy event in the
vicinity of the Mount Crawford granite gneiss.
There ure a few exposures in which measure-
ments made On late fold siructures suggests
anomalous unentations. for example, the irre-
gular syniorm in the south-west, These
anomalies muy be related to a conjugate P;°
arian Fy event,
The Mount Crawford granite gneiss replac-
ing the eastern part of the pelitic sequence
contains a pronounced schistosily with the
299
charagtenstic gneissic S: microstructure of the
pelitic schists including the characteristic im-
bricate stacking arrangement of individuil
mica plates within the mica trains. A strong
lineation, considered to be L(S)-S.), bies within
the schistosily and is defined by elongate mica
plates and quartz and felspar grains, The
schistosity has a mean strike of 349° and a
dip o! 56° east and the lineation plunges 33”
to 149° (see Fig. 6, sub-area G), The unitonn
orientation of the L{S,-S.) lineation t related
to Lhe uniform initial orientation of S,; and 5S.
and the fact that the granile gneiss has largely
escaped the F, deformation event. The granile
goeiss would appear to have behaved aos a
solid block after the main metamorphic. peak
was teached early in the Ps eveni. As a con
sequence it has escaped the lale or post-F
relrogressive event and the effects of the Fy
deformation. Skialiths of pelitic schists within
the granite gneiss have becn affected by the
F, and Fy events, but not the F, event,
Numerous amphibolite dykes of basaltic or
doleritic parentage have been mapped in this
area (Mills 1963), (hey ate confined to the
coarse-grained schist sequence east of the
Warren Inlier and also penelrale the granite
gneiss. They have a distinctive microstructure
made up of a schistosity and strong lmeation
defined by hornblende prisms. Within dykes
cutting the granite gneiss these slructures are
concordant with the S.-L{S,-S.) microstructure
of the enerss. Within the pelitic schist sequence
both Fy and F, falds have been observed
a
lig. 4. Profiles of F: folds in Adeluidean succession. ;
uw. and 6—Moderately light Fy folds in metasandstone. Note Ss crenulation cleavage deforming
earlier 8 schistosity in mica-rich layer in a, and weak So axial plane structure defined by flattened
quartz grains and ortented mica plates in &. Traced trom photographs, Location 917126.
c—Tith! Fo folds in quartzofelspathic layers (stippled) in micaceous gneiss of the thick pelitic
sequence, Nate thut an early schistosily ($1) preserved in the quartzofelspathic layers patallel
in the layering is folded in the Fy deformation. In the micaccous gneiss the 5. crenulation
cleavage has been Jargely transposed into a new schistosity, Traced from photograph, Location
28116,
d— FE. folds in quartz-felspar-sillimauite-mica gneiss of the thick pelitic sequence, An early S,
schistosity, apparently parallel to the prim
ary compositional layering, is almost obliterated by
ihe imposition of the S, cleayage axial plane to the folds. Traced from photograph. Location
92ST.
¢, and {Tight F. folds in coarse-grained mica schist containing quanizofelspathic layers (quartz
stippled. inica shaded). An earliet schistoslty (S,), outlined by mica plates, is well preserved in
both mica-tich and quartz-rich layers (S.). Traced from photomicrographs. Location 934094.
ge—Tracing from photomicrograph showing early stage in the development of Sy mica-rich
layers during the process of crenulation of earlier schistosity (S.) im a quartz-biotite-sillimanite
achist enclave in the granite gneiss, In the granite gneiss the S, structure 1s largely desiroyed but
ihe Se gneissasity commonly preserves the characteristic echelon or imbricale arrangement of
mica plates. Location 939102,
A—E: folds in fine-grained quartzofelspathic bedding laminations (stippled) in doloritic
phyllite. The axial surface structure in the phyllite is a very fine crenulation cleavage (shaded),
Wote the cross-cutting quartz veinlets which were ptygmatically folded in the Fz deformation,
Biolite zone phyllites west of Williummstown-Meadows Favtr-
Location 890120.
Traced from phoiomierograph,
)
— AF
Sey (gs
—_——_
a a
(Miah
(TY
yates iy
i aE Hi Uf!
pbs
fo" tf!
Yess
eens
K. J. MILLS
ed)
y; yy
oO i /
if}
3] t/
Af
300
he
Fig. 5
STRUCTURAL GEOLOGY OF WARREN NATIONAL PARK
within the amphibolite dykes. Some Fy folds
involving amphibolites have been recently
exposed on the south side of the reservoir in
the cutting for the road leading Lo the new
bridge. The amphibolite: were apparently in-
truded during or prior to the F, event.
Veins and pods of pegmatite and quartz
are intruded extensively through the pelitic
sequence, Some of these were strongly sheared
prior io the F, event and were subsequently
folded in the Fy and Fy events, but most of
the coarse-grained massive varieties appear to
have been intruded after the Fu event, Some
of the smaller pegmatite veins were folded
during the F, event. Some large tourmuline
porphyroblasts were observed to have grown
across Fy ‘structures,
The kyanite-andainsite zone
Medium-grained biotite-quartz-felspar
schists, marbles and cale-siheate rocks belong-
ing to this zone are exposed cast and south
of the Murray Vale Thrust and between the
Ukaparinga and Williamstown-Meadows faults.
in the north-west. The schists in this sequence
ure characterised by a simple planar schis-
tosity, S;, made up of oriented mica plates
which ate usually evenly dispersed through the
rock, This schistosity is almost parallel to
bedding laminations in most exposures and no
mesoscopic er macroscopic F, folds have been
found. Deformation structures Telated to the
F,.and Fy events are rarely observed und are
only weakly developed. A notable coarsening
of grain size, especially of mica plates in mica
rich laminae. and the growth of post-tectonic
muscoyiie porphyroblasts occurs low in the
It
sequence udjacent to the higher grade block
and near pegmatite intrusions. This could be
due to some kate contact heating aller thrust
sheet emplacement. Peymiutite Intrusions of a
massive or zoned type are rare and are con
fined to a few veins running parallel to S.
immediately east of the Murray Vale Thrust,
No amphibolite intrusions are known to cut
rocks of this zone,
Bedding, defined by compositional variation,
is well preserved on all scales in this sequence
and where the exposure is sufficient the beds
are found ta have great lateral continuity.
Stratigraphic facing indicators are rarely
observed, but the sequeuce is considered to
dip east and to face east in the east and to
dip west and face west in the north-west
block, the two wreas of exposure representing
the limbs of a very large F, anticlinal struc-
ture. The schistosity S,, assumed to be axial
surface to the mucroscopic F, structure, is
observed to lie at a smal! angle to Sy, usually
less than 10° in the more micaceous rocks.
This tends to promote splitting along ihe
bedding surtaces of the quartz-felspar-biotite
schists to yield slabby blocks which were ap-
parently favoured for building purposes in the
early days of seitlement. Microscopically the
5, surface in the schists is defined by the
orientation arrangement of biotite plates,
which tend to have an even spatial distribution
except in strongly knotted schists. where cus-
pate concentrations of biotite plares have
grown at the margins of porphyroblasts. ‘These
porphyroblasts are mostly retcogressed to fine
sericite but contain rare telics of andalusite
und kyanite. The size of inclusions in the
a
Fig. 5. Profiles of F, folds in the thick pelitiy sequence of the Adclaidean succession,
a—Open Fs folds in mica schist, The main schistosity (S:) is apparently parallel to the com
asitignal layering. Sketch from photograph. Location 928117,
_—Polyclinal Fy folds in mica schist. Main schistosity ix Se Fs fold outlined by quartz fens
has been folded in the Fy deformation. Traced from photograph. Location 926116.
c—Open Fy folds in micaccous metasandstone, Main folintion is S:. Traced from photograph,
Loeation 924109.
d—Open F- folds in miva schist. Main schistosity is S. which appeats to he parallel to the
quartzofelspathle fayer at the baxe of the diagram (stipplei)-
Traced [rom photograph, Location 926117,
Note cross-cutting quartz lens,
e—Tigm F: folds in quartz-felspar-mica gneiss. Main schistasilty is Ss Traced from photo-
micrograph, Locution 929119,
{—Disharmonic Fy folds in mica-quartz gneiss (quartz stippled, mica shaded). Main schistosily
is Ss, Note late cross-cutting fault. Traced from photomicrograph. Location 918060.
v—tTieht F, folds in quiarlz-muscovile gnetsy showing incipient development of Sy crenulation
vleavaze (quartz stippled, mica shaded).Main schistosity i5 S:. Traced from photomicrograpls,
Location 943118.
* Owing to differences in the structural development in the high and low-grade sequences thecorrelalion
of mesoscopic structures is p mutter of conjecture, The author has adopted the simplest correla-
tion campatible with his present knowledge of the area.
302 K. J. MILLS
SUBAREA E 5
| 35. 4 LIBS: 4
Soc
2 Lal?
= L Lge
ere ARES F
at
a Se
‘ “—
{
SUB- aes G
251 Sz
i vine)
[Max 6Bur
198 LSe
imax da-laa; Se
| - SUBAREA, H
ul _—_
!
i
WA
ao,
SUBAREA A (BASEMENT)
58 SCHISTOSITY 6 MINERAL STREAKING | 25 AXIAL PLANE 32 CRENULATION
PLANES = LINEATIONS o CRENULATION po, LINEATIONS «=
CLEAVAGES * a mA
Fig. 6.
STRUCTURAL GEOLOGY OF WARREN NATIONAL PARK 303
Soe S,+ Lipa “ S:: Lisig2
Leis»
SUB-AREA aw
1 Lips
SUB-AREA K
4 LIBIS: | 15 So 1 Ss
la LE Nop |
SUB-AREA L
11s, F 87 S {7 LB) 18 Sy
7 A nape 2 3 Lil: a4 LieySs
SUBAREA? « MN
\ Lyeyg2
SUBAREA: s OPQ adh
13 Le oe 2 LIBlgs |
Fig. 6, Lower hemisphere equal area projections of structural elements measured in all sub-areas shawn
in the inset figure except sub-area R, True north at top of diagrams. Contours at 0.5, 5, 10, 20,
30% per 1% area.
304 K. J, MILLS
:
Na ees
RESERVOIR
DMO Ih
FINE-GRAINED
MICA SCHISTS
|
METASANDST ONE
bad WARREN
INLIER
Ss > Se
SUB- a, * LB)! a S, LiBIee a
Ss
AREA L(B) 3? *
xe ‘Sh Sz
35, pot i L(Big! | 64 Sz 2 L(ahge
F2-072)
Rq
34 LIB)SS
(MAX GO-O701 42
L.
/ ;
R {
| A ae 25 Lip) Ss 33 LBS
: (Max. ¢5- 058) z
72S, 172 3;
(MAK (wax,
fa— 2a)
Ro te
a
2
109 .(B)$
| \MaAwe at 's
L
Fig. 7. Lower hemisphere equal area projections of structural elements measured in coatse-grained
Adelaidean schists of sub-arca RK. True north at top of diagrams, Contours at 0.5, 5, 10, 20.
30% per 1% area.
STRUCTURAL GEOLOGY OF WARREN NATIONAL
porphyroblasts jo relation to matrix grain size
indicates gerowlh during and after the F,
Ueformation event. In other rocks ellipsoidal
quartz, felspar and carbonate crystals, and the
Jong axes of amphibole prisms, may define S,.
A moderale quartz c-axis fabric has been noted
im some specimens of quartzofelspathic schist
and ay early quartz veinlet cutting the S,
surface of a quartz-felspar rich schist in the
north-west block was observed to be recrystal-
lised and to have uw strong quartz fabric in
which most c-axes were aligned within the S,
surtace,
A fait to strong mineral streaking lineation
is notable within the S, surfaces cof some
exposures. This lineation is defined by elongate
mica plates and porphyroblastic knots in
schists and by amphibole. prisms in calcareous
rocks. A set of cross joints is commonly scen
lo lic perpendicular to this lineation. In the
mica schists the lineation. is defined by a ¢om-
binalion of two cllects, Firstly the mica plates
are up to twi¢e as wide parallel to (001) in
the direction of the lineation than they are
across it, the biotite plates being considerably
more stumpy in form than assoctated musco-
vite plates. Secondly the mica plates 2re much
more diversely oricnted in sections cut per-
pendicular to the lineation than they are in
sections cut parallel to it: that is, the mica
plates are fautozonally arranged about the
lineation axis. In the absence of suitable strain
markers in the present arca it is not possible to
determine whether this lineation as due to the
mimetic growth of crystals in lines of bedding-
cleavage mtersection, L{S,-S;}, of is a tectome
strain lineation, L(S,), developed during meta-
morphic mineral growth, The penetrative
nature of this lineation in many cocks, and
strain shadow ejfects around syntectonic por-
phyroblasts, favours the latter.
No mesoscopic B(Sy+S,) folds have been
observed, In sone schists in the north-west
block o mild crenulation in the S$, surface,
showing syntectonic crystallisation of the mica
plates in the crenulation hinges, suggests
effects of the F, event. Some mesoscopic
F. folds of hand specimen size have been
found as Aoat in the north-west block, but they
are apparently quite rare. These folds show §,,
parallel to 8, folded into V-shaped sivles with
planar limbs, well defined hinge regions wad
thickened noses.
Likewise the effects of the Fy, event are
weakly developed in this sequence, The
PARK 3s
Gumeracha Goldfields antiform in the south-
east has macroscopically flexed the 8, and 8,
surfaces, but otherwise, only aecusional bax-
like post-crystalline c¢renulatians, showing
strong strain effects in the minerals of their
hinge regions, are considered to belong to the
F.. event.
Biatite zone
West of the Williamstown-Meadows fault
line fine-grained impure dolomites, dolomitic
phyllites, phyllites and phyllitic slates bhelong-
ing to the biotite zone are also characterised
by multiple defarmation, Evidence for ihree
overprinted deformation phases can be recog-
nised in some exposures and provisionally
equated wath the B,, Fy and Fy events. These
rocks have a very fine grain size, quartz and
micas averaging 0.01-0.015 mm, and invari-
ably have a strong §; slaty cleavage labric
defined by the shapes of curbonate grains and
the parallel orientation of muscovite and bio-
lite plates. "Strain shadows" are presence
around some opaque minerals, The ¢c-axes of
quurle grains possess a strong orientation in
the S, fabric of some phyllites, with most
c-uses tending to Te im the S, surface.
S; is seen to lie at large angles to S, in some
exposures and broad hinge regions of B(S,-5,)
folds are present. So is usually well defined by
compositional layering or Jaimination.
At 890120 well developed finely spaced
strain-slip or crenulation cleavage, Sy. was
observed in phyllite. Microlithons between the
crenulation cleavage planes. are about 0.3 mm
in width. The S, cleavage within the micro-
lithons as bent into the S, crenulation cleavage
surfaces and this process ts ackompanied by
unm increase i mica content within the Sy,
surfaces. Apparent displacement on the S, sur-
faces crossing the phyllite is not represented
as displacement in cross-cutting quartz vein+
lets, but as. sharp monoclinal folds with
atienuated limbs crossing the S. surface,
Where S, is developed, mesoscopic 8(S)-S.)
folds are observed in the bedding (Fig. 4h).
QQuartzose layers are buckle folded, while car-
bonate-rich laminated phyllite layers are strain
slipped. ‘The folds are disharmonic on all
ecales, but the So surtuces tend to have 4
consant orientation. Thin quartz stringers
were seen to have been ptygmatically folded
un a microsenpte scule in the Fy event. Later
more brittle Kink folds are rarely observed
and might be attributable to a weak Fy event.
Oe
FAULY SPRUCTURES
Thrust faults
Several thrust sheets have been mapped in
(he present area on the basis of metamorphic
and structural discontinuities. Vhese thrust
sheets Wert emplaced during or after the Fy
event and were subsequently folded in the F,
eveot. Four principal thrusts are named. The
Wirriynda “Uhrust, passing near “Wirrianda’
homestead situated on the southern shore of
the reservoir; the Watts Gully Thrust, passing
close to the site of the richest gold discoveries
in the Gumeracha Goldfields; the Sanctuary
Vhrust, largely confined to the flora and fauna
sanctuary in the south-west; and the Murray
Vale Vhrust, named after the pastoral property
south-east of the headquarters of the Mount
Crawtord Stule Forest. The Jast three named
faulis appear to be hinged near the southern
margin of the mupped area. The opalised
Iremolitic rock bed at the contact of the
mictasandstone and the course-grained schist
sequence has proved to be a useful marker
horizon in the identification of sequence repe-
tiliom in the thrust sheets, There are no proven
natural expasures of the thrust Fault surfaces
in the area, and critical thrust shyet interplay,
particularly in the south-west. ig obscured by
alluvium and poor ouicrop, There is some
evidence From the study of the mesascopic
orientation data that the higher sheets rotated
clockwise oVer the lower sheets in the south-
west corner of the area. This may be the
result of rotation during the thrusting move-
ments or differential rotation of the sheets in
the subsequent F, event. Numerous exposures
of sheared and mylonitic gneisses in the thick
pelitic seyuenve are thought Io have been deve-
loped during the thrusting movements,
On the northern slopes af the reservoir the
Wirrianda Thrust is marked by a sharp
contact between metasandstone and course-
grained micaceous eneisses. South of the
reservoir the sandstone cuts out and the hase-
ment gneisses are brought into contact with
the schists and gneisses of the Adelaitlean
sequence, Although exposure is fairly good
here. similaritics between rock types and the
overprinting of mesoscupic Fy folds From the
Adelaidean schists into the basement racks
have obscured the actual fault surface, Further
south the upper section of the metasandstone
\nit reappears. but the fault zone is everywhere
obscured by soil cover, North of Watts Gully
(he fault line follaws the bottom of a steep
Veshuped valley and its actual path across the
K. J, MILLS
mctasandstone further ta the south-west has
not been identified. but it is assuined to
eventually rin mto the Willramstowm-Meadows
Paull.
Structural evidence jin support of the Wutts
Gully ‘Thrust is obtained from an area of
woderate exposure near the centre of the
patch of virgin scrubland of the flora and
fauna reserve south-west of Dead Horse Gully,
Here considerable discordance exists between
structures on either side of the mapped fault
trace, and 4 narrow splinter of pebbly
micaccous mctasandstone appeurs ta lic
within the fault zone, The Watts Gully Thrust
is also responsible for a repetition of the strati-
graphic sequence, Near Watts Gully the fault
trace can be followed us 4 junction between
schists and metasandstone and js marked in
places by a distinctive white albite rich rock,
which may carry tale or actinolite crystals.
Under the microscope this rack is largely cam-
posed of well-crystallised untwinned plagioclase
with a composition near pure albite, carrying
occasional rutile inclusions, Some interstitial
quartz grains with slightly undulase extinction
are dispersed through the albite, Scattered
muscovite plates, preserving & microstructure
like that of Sy in the pelitic schists, ure
strongly bent and are encased or adjoined by
the unstrained post-tectonic albite.
The Sanctuary Thrust is of limited extent
and is defined on the basis of a repetition of
& belt of metasandstone shove pelitic schists,
The fault trace is also marked in a few places
by a peculiar white albilte rich rock similar
to that described above,
Movements on the Murray Vale Thrust
were of greater magnitude than those on the
underlying thrusts and resulted in the conjoin-
ing of racks of markedly differing metamorphic
and structural character, as described in
previous sections, Fxposures of rocks near
the thrust surface are very rare, and for
most of ils length the thrust line is marked
by a conspicuous strip of soil cover. In the
field the position of the thrust has been placed
on the basis of the readily mapped distinction
between the coarsely crystalline rocks of the
sillimaniteanuscovité zone and the medium cto
fine-grained schists of the andabusite-kyanite
zone. Much of the eastern margin of the
granite gneiss has been reinlerpreted as the
thrust surface. The banks of a road cutting
on the improved section of the mitin road
south of the reservoir, about half a kilometre
north of the headquarters of the Mt Crawford
STRUCTURAL GEOLOGY OF WARREN NATIONAL PARK
Stale Forest, expose rocks on either sid¢ of the
Murray Vale Thrust, Fine-grained schists and
the thick opalised tremolite rock bed lit in
conjunction with weathered amphibohle wand
coarse-grained mica schists with intrusions. af
pegmatite, The faulted contact is marked by
& Narrow zone of green clay-like pug.
The Murray Vale fault linc has been super-
ficially traced north of the reservoir where it
is marked in a few places by a sugury while
albite rock containing green actinolilic spots.
After passing through a left-handed displace-
ment, probably a large F, fold, it traverses
the centre of the large quarry excavated for
clay und sillimanite. Near the base of the
quarry the fault zone is seen to be less than
half a metre in width and filled with pale
greenish clay-like pug. The fault surface dips
easterly at about 70° with some irregularities
caused by open Fy folds, On either side of
the faull extensive shearing, especially in the
higher grade rocks to the west, has resulied
in the formatiin of the unique Williamstown
damourite schists containing kyanite uni
corundum, massive sillimanite pods and clay
Uepusits. The clay Ueposits seem to be related
to Inte stage alleration within the faule zune.
Wesl of the fault ale a variety of high grade
poeises Showing Jess allteralion to clay away
from the fault surface. Fine-grained lower
grade schists are exposed cast of the fault.
Same are very fine-grained pyritic schists
which are extensively altered to clay near the
fault, although still preserving Hresh pyrite
cubes near the base of the quarry. Some open
fokis, presumably of the F,, event, are visible
in the lower grade rocks. North of the quarry
the Murray Yale Thrust probably passes
through another lurge F., fold resulting in the
uppearance of fine-grained quartz-felspar-
biotite schists displaying open shallow north-
phinging F, foldy in small quarrics north of
the forked junction of the rounds Jeading to
“Spritigheld"’ and the Australian Industrial
Minerals quarry.
The Ukeaparinga Fault
The Ukaparinga copper prospect in the
north-west corner of the area is situated
within the brecciated zone of a major meta-
morphic and structural discordance, herein
named the Ukaparinga Fault, Displacement on
this structure must be at least as great as thot
on the Murray Viele Thovist, but it is ancertain
as to whether this faull should be grouped
with the pre-Fy thrusts or i a brauch of the
V7
nearby post-F, Williamstown-Meadows Fault,
which it joins a little south of the South Para
River, A fillle 19 the west of the Ukaputinga
Fault, # parallel fault line has been traced on
ihe grownd in an area of moderale exposure
of the sehisty of the kyunite-andalusite zone
sequence, but its importance as a fault struc:
lure has not been determined.
The Williamstawn-Meadows Fault
An important fault line, separating fine-
grained biotite zone rocks from coarse-grained
amphibulile fucies rocks, can be traced in a
north-south direction on the western side of
the area, In eurlier literature this fault his
been referced to as the Kitchener Fault, but
as it appears that this name is based on a
misconception, and since the author has been
able to follow the path of this fault as a con-
tinuous line from well north of Williamstown
to well south of Meadows, the name Williams-
town-Meadows Fault is preferred,
Within the present area the actual fault zone,
along most of its length, appears to be very
narrow and sbhurply defined, although some
exposures of tectonically emplaced foreign
biogks lying within the fault zone fave heen
shown on the map as fault melange. Where
the fault line crosses deeply incised valleys, a
sleep easterly dip of about 60-70" has been
estimated for the fault surface at several
points. ‘This would suggest that the fault is
of the steep reverse Lype, bul unlike the thrusts
described earlier, movement occurred after the
F, folding episade,
A uarrow zone of brecetated phyllite marks
the fault zone al ong exposure north of the
South Para River. South of the river fine-
grained dolomites are brought against the fault
surface im several places and close to the
fault these rocks have been recrystallised to
medium-grained white dolomilic marbles can-
taining large plates of muscovite and talc and
minor amounts of quartz and felspar grano-
blusis and an opaque accessory, The ¢equi-
granular hornfelsic texture and the partially
dissolved nature of twin relics within the
carbonates suggesis That these marbles were
uffected by some form of contact metamar-
phism, It seems that the higher grade hanging
wall block was still sufficiently warm during
and after the fault displacement to contact
metamorphose the impure dolomites near the
inmediate contact with the fault surface.
In the south-western corner of the Warren
National Park float and outcrop of felspathic
Is
and micacedus schists and gneisses of foreipn
texiural appearance, opalised tremolitic
marbles and metasandstanes testify to the
presence of a mélange lens within the fault
zone A further lens containing peculiar mica-
tich schists defines the fault zone separating
metasandstones from dolomitic phyllites
almost half way between this locality and the
South Para River. The microstructural features
of the schist blacks within the fault zone can
be provisionally correlated with the 8), 5,
and S. structures of the area, The meta-
morphic wrade of these schists seems (o be
that of the upper greenschist or Jower amphi-
bolle Caciex, A small stauroliie crystal was
found in a specimen of bjotite-rich sehist.
Apart from some kinking and bending of mica
plates and quartz grains and the introduction
of thi stringers of potash felspar, the fault
movenjents appear to haye had little effect on
the structure of these schists, and the faulting
appears to have occurred when the metas-
morphic temperatures were too low to permit
any significant recrystallisation.
The amount of displacement on this fault
must be very large judging [rom the meta.
morphic diflerences between rocks on either
side of the fault surface. Miles (1950) suz-
gested a minimum displacement of 5,000 with
a probable displacement of ahout 10,000’, Dis-
placement of the order of 3 to 5 thousand
metres does not seen incongruous, but a better
estimate of the amount of displacement must
uwail a amore detailed study of the strati-
Braphic and metamorphic relationships along
mast of its length,
burlier workers (Miles 1950; Sprige 1945)
have considered the possibility of Tertiary to
Recent movements on this fault, the evidence
for this being based on the marked fault scarp
expression and the occurrence of erosional
surfaves and gravel beds of supposed Tertiary
uge west of the fault line near Williamstown.
However, hills composed of Adelaidean bed-
fack in the south-west of the present area are
al the samme height on either side of the fault
line, and there is no evidence of recéni re-
juvenation on the fault surface. The fault
acirp i$ believed to be due to the erosive
power of the Soath Para River and its tibu-
tariey removing the more readily disintegrated
biotite zane: phyllites and dolomites west of
the tault, after crossing the resistant Warren
Infier and iis mantle of metasandstones and
schists. This erusional scarp was probibly in
existence during the formation of the Tertiary
KF MINS
gravels and lateritic surfaces near Williams-
town, with the Warren Inher and its swirround-
ing mantles of high-grade rocks protruding
above the Tertizry erosion levels,
CoONELUSIONS
(£1) Basement-Adeluldean relationships
Howchin (1906, 1924) clearly recognised
the existence of basement rocks in the porge
of the South Para River, south of Williams-
lown, and claimed that an unconformity
separated the basement from the underlying
metasandstones of Aldgate Sandstone type,
Hossfeld (1935) claimed to have recognised
an actual exposure of the unconformity near
the Warren Reseryoir weir, but this could not
be confirmed in the present study, Luter
workers (Campana 1953: Mills 1963) tailed
to recognise a basement inlicr. Campana
apparently regarded the lowest schists a5 metu-
somatic alteration products of the Aldgate
Sandstone equivalent and Mills interpreted the
schists below the metasandstone as an early
Adclaidean pelitic sequence, The present study
hus confirmed Howvhin’s claim for the
exisfence oF a basement inlier, Marked struc-
tural discontinuity across the observed uncon-
formity surface discounts the possibility of an
early Adelaidean pelitic sequence. The cosrse-
grained schists and gneisses above the mceta-
sandstone unit Jithologically resemble the
basement yneuses, but they contain mesuscupic
structures which may be correlated with unu-
logous. structures. in the metasandstone unit
and not with structures in the basement
gneisses.
Io the nearby well-established Houghton
Inlier, Spry (1951) showed thal schistosities in
the basement and in the Adelaidean mantle
were generally parallel. ‘The schistosity in the
basement was interpreted as a retrograde
schistosity which was associated with the
Pulteozvic folding of the Adcluidean succes-
sion and updoming of the basement, although
tere was some evidence (Spry 1951, p. 120)
for shearing of the basement prior to Ade«
dsidean sedimentation, Campana (1955) sug-
gested that movements in ihe basement of the
Adelaide region were responsible for the
folding of the Adelawewn sedimentary mantle.
the lower beds of which were consequently
adjusted into tight folls characterised by
sharply overturned anticlines, with attenuated
limbs grading into overthrusis and broad
¢ortugated synclings, Talbot (1962) curried out
an extensive study of the southern part of
SIRUCTURAL CEOLOGY OF WARREN NATIONAL PARK 49
Fig, ¥,
Hee alts
fa a if shee)
Block diagram constricted to scale in orthographic projection on a line of sight plunging 35°
towards 253° showing the forms of the principal stractural surfaces in the vicinity of the Warren
Reservoir, The structural surfaces shown are the thrusts and faults (cross-hatched), unconformity
-etween the basement rocks of the Warren Inlier and the Adelaidean sequence and the lop
of the Aldgare Sandstone equivalent (stippled). The surface of the block corresponds to the
high water level in the Watren Reservoir but detailed adjustment to the map pattern to allow
For topographic variitions have not been made, The F, synform outlined by the unconformity
in the centre of the diagram is overprinied by macroscopic be folds, and other Fu folds are seen
near the Warren Reservyir, The structural surfaces are folded by larze Fs folds in the southern
part of the diagram «nd lesser Fz folds are interpreted to deform the Murray Vale Throst: lo
the north of the reservoir.
the Houghton Inlier and its Adelaidean mantle
abd considered that the chlorite grade base-
ment “acted a5 a simple core structure and
adjusted to the folding of the overlying
Torrens Group" by passive movement along
foliation planes produced in a pre-Adclaidean
Tetrogressive phase.
The present study has established that the
lower portion of the Adelaidean succession
is welded onta the basement rocks of the
Warten Iplier ond that the basement rocks
were rejuvenated during the deformation
events and the extensive metamorphic recrys-
failisation which affected the overlying Ade-
laidean sediments during the lower Palacovoic
Delaincrian ofogeny. The surface of uncon-
formily has been infolded into the basement
encisses and there is little evidence of any
differential mayement at the boundary between
the basement rocks and the overlying sedi-
mentury mantle. The basement rocks of the
Warren Inlier have undergone 4 partial tee-
tonic reconstitution ji which the lower
Palaeozoic folds wens imposed on the older
pre-Adelaidean structures.
(2) Folding of the Adelaidean succession
Regional studies in the Mt. Lofty-Olary
orogenic arc have indicated that the upper
Proterozoic (Adeluidean) and lower Palaeozoic
successions were affected by only a lew gentle
epeirogenic episodes (Sturtian tectonisin, Dut-
tonian folding, Cuassiniaw and Wailpingan
movements) prior lo a major orogenic revolu-
tion. (the Delamerian Orogeny) in lower Ordo-
vician time (Thomson, in Parkin 1969), This
ofogenic cycle brought about cxtensive fold
deformation, created a helt of low to: inter
mediate pressure metamorphism and’ granite
intrusion, and brought an end to geosynctinal
sedimentation in the Mt. Lofty Range region.
The syntectonic Palmer granite, intruded into
the core of the metamorphic belt in the eastern
Mt. Lofty Ranges, bas been isotopically dated
at 490 million years (White, Compston &
Kiceman 1967).
Detailed structural studies in several key
areas in the Mi, Lofty Ranges has established
that three important episodes of fold deforma-
tion look place at various stages during the
‘ia K. J, MULLS
metamorphism, Oller & Fleming (1968) syn-
thesized previously published and unpublished
work relating to the deformational and meta-
rourphic history of the Mf, Lofty Ranges and
ussumed that euch of the three folding
episodes (F\, Fa and F,) were synchronously
developed across the whole deformed belt, but
it seems that more key areas will need to be
unulysed before this premise can be verified
and a satisfactory unravelling of the deforma-
tional history is achieved.
The present structural study of the upper
Prolerozoic rocks around the Warren Inlier
has established that three distinct tectonic
episodes myolving fold deformation have
occurred in succession in relation to meta-
morphic recrystallisation,
F, deformation
Bedding surfaces (Sy) of the Adelaidean
succession Were deformed into large regional
folds during recrystallisation of the sediments
under conditions of rising metamorphic tem-
peratures and mica plates and other meta-
morphic minefals. grown or deformed syntec-
tonicully, developed 9 strong axial surface
schistosity fabric (S,) containing an inter-
related axis of elongation, 1{S,), which is
expressed aS a minecal streaking lineation in
some exposures. The commonly observed near
parallelism of S, and S, suggests that these
early folds possessed an isoclinal style with
extensive oear planar limb areas. Mesoscopic
F, folds ate apparently rare and difficult to
identity, and litthe can be deduced Irom the
initial geometry of the F, deformation within
the vonfines of the small area studied. Struc-
tural analysis of the bedding attitudes in the
basul iietasandstone suggested that the hinwe
lincs of macroscopic F, folds were gently
plunging or subhorizontal and it can be
assumed from regional studies elsewhere in the
ranwes that the hinge surfaces of the F, folds
were dipping stccply cast and striking north-
south purullel tu the tread of the orogenic belt
(Offler & Fleming 1968), The regional struc-
ture which Campana ¢? af. (1955) named The
(Lookout Tower overturned anticline can be
considered to refer io the demal structure
comprising the culmination of the Warren
Inlicr, although this culminution is the cam-
bined resultant of the Fy, Fy, and F, folding
events,
‘Textural-mincralogical evidence sugecsts that
metamorphic lemperateres hid reached a peak
by the emf of the F, deformation, or during
the interkinematic period between Lhe Fy and
F, events and the marked metamorphic zona-
tion observed in this area Was impressed al this
time. In the higher grade schists the mica and
quartz grains defining the plane-linear S)-L( $1)
labric had developed a coarse praili-size and
sillimanite fibres and prisms had grown within
the $, surfaces prior to the commencement of
the Fo deformation. Despite high-grade mets-
morphism sedimentary clasts and textures are
slill preserved in must specimens. of the coarser
arenites, even in specimens exhibiting @ sirong
S,-L(8,) fabric. In those metasediments
assumed from their composition to have in-
iiated as pelites and semi-peliies recrystallisa-
tion and grain growth has obliterated all signs
of sedimentary structures in the coarser
grained rocks of the sillimanne-muscovile
zane, but thin bedding laminations are pre-
served in the finer grained schists of she
kyanite-andalusite and biotite zones,
F, deformation
This dclormation episode commenced at or
near the peak of metamorphic tenyperature,
allowing recryslallisalion and grain growth
processex to become dominant over strain
preservation in some rocks, and allowing the
F, strain effects to outlest grain growth in
others. The resus of ihe F; deformation are
expressed in two principal forms:
(1) The development of intensely appressed
folds in the earlier Sy and 5S, surfaces and the
imposition of a strong crenulation cleavage-
schistosily (Sy) with a north-south strike ane
a steep easterly dip, and
(2) the development of major late-meta-
morphic thrusts which now divide the area up
infe several distinct stryotural-metamorphic
fumes.
The penctrative effects of the F., deforma-
lion are most pronounced in the coarse micu-
rich schists and gneisses of the sillimanite-
muscovite zone where intensely deformed isu-
clinal folds in lithological layering have 32
well-developed crenulation cleavage (So}
(crenulation of the S, schistosity) grading into
a new coarse-grained schistosity as yn axial
surface structure. In many exposures the Sy
schislosity has become the dominant penetra-
tive structure and traces of the former §,
schistosity are preserved only in the form of
a strong mineral streaking Jineation L(S,-So).
The Mount Crawford pranite gneiss formed
during the F. deformation and presetves a
well-developed S. schistosity and a constantly
oriented mineral lineation interpreted as the
STRUCTURAL GEOLOGY OF WARREN NATIONAL
L(S,-S.) intersection lineation, Basic dykes,
intruded helore or during the F, event. re-
crystallised to schistose amphibolites preserving
a strong amphibole prism lineation, which is
interpreted as the result of mimetic growth of
hornblende prisms parallel to the .(S,-Sol
intersection lineation,
The effects of the F. deformation are Jess
pronounced in the mutasandsiones. and,
although F, crenulations are well developed m
the more mica-rich Varieties, intergranular
movements were probably daminant, Meso-
scopic F, folds in the metasandstones show
strong appression und Hattcning of the quariz
grains info the S, surface. In some ufcus some
of the plastic strain induced im the areniles
during the F, deformation outlasted recrys-
tallisation and strain relief. leading to the
developrnent of well-orientated quartz c-axis
fabrics.
In the lower grade kyanite-andalusite and
biotite zone blocks the FP. deformation is aot
50 apparent and the S». crenulation cleavage
is only locally evident, although some local Fy
mesoscopic folds are to be found and show
a similar tight uppreasion.
The Fy, deformation culminated in the ap-
pearance of several steep thrusts and mylo-
nilic zones iwvolving considerable, but as yet
unknown, amounts of displacement,
Following the Fy deformation, a sceond
static interkinematic period is envisaged in
which the strain preservation fromm the Fs
deformation became relieved under greenschist
fucivs conditions resulting m the exilensrve
retrouression of higher grade minerals to
quartz, albite, chlorite, sericite and rutile, The
sercite has undergone patchy regrowth to
larger randomly oriented muscovite crystals,
The growth fabrics of these late stage minerals
tend to be isotropic except where they were
mimetrically controlled.
Fy deformtation
Most parts of the area covered in the
preseril survey were affected hy a rather more
brittle deformation late in the metamorphic
Nistory resulting in smucrescopic and meso-
scopic. folds in earlier structural surfaces. The
Fk, deformulion is more pronounced in certain
zones. F, folds are rounded te angular and
PAKK 311
show varying degrecs of appression. ranging
From open warps with eust-west hinge surface
trends to tight folds with north-south hinge
surface trends and a weakly developed crenus
Jation cleavage (S,). Both the hinge surfaces
and the hinge lines show considerable varia-
tion in attitude, Minerals grown dunng the
earlier relrogressive metamorphism became
deformed during F, folding and show signs
of both plastic and brittle strain. The preserva-
tion of such fime F, strain features as quartz
deformation lamellae suggests What very little
erystal recovery took place after the F, event.
The fold deformational history outlined
above is in general accord with the conclu-
sions of other structural studies carried out in
the Mt. Lofty Ranges (Offler & Fleming
1968). At this stage it seems valid to correlate
the present results wilh those from the Pewsey
Vale area 10 km north-east of the Warren
Reservoir where Offler (1966)! found three
deformational events in the Adelaidean sue-
cession which appear analogous to those
described here. In the Pewsey Vale area F,
tolds were rare and S, was almost parallel
to Sy, some mesoscopic F., folds were deve-
loped and open macroscopic Py felds dami-
nated the regional structure. Metamorphism
had reached its peak late in F; or during F,
and was waning during the FP, event, Strong
crossed girdle quartz c-axis patterns developed
within quartzites after the F, phase but priot
to the Fy event. In the adjacent Kanmantoo
Group sediments at Pewsey Vale, Offler re-
corded two deformational cvents which he
interpreted as F, und Fy, with metamorphism
oullasting the F, event, In the Cambrai-Spring-
ton region Mills (1964)° found that the Kan-
mantoo Group metasediments were tightly
folded in a dominant early deformational
event with focal over'priating by a second fold
event with the metamorphic peak being
reathed during and folluwing the second
event,
(3) Faulting
Two varieties of faults have been distil
guished in the present survey), firstly, faults
and mylonile zones considered to be of a high
angle thrust type, which developed late in the
F, deformational event and wert subsequently
folded in the F. event: secondly. a high angle
* Offer, R, (1%66)—The stenctuve and metamorphism of (he Pewsey Vale atea, north-east of Williams-
_ town, Soulh Ausiralia. Univ. Adelaide, Ph.D. Thesis. :
* Mills. K. J. (1964)—The sirucrural petrology of au area east of Springton, Sout Australia, Univ
Adelaide. Ph.D. Thesis.
312
reverse fault, the Williamstown-Mcadows
fault, which developed after the Fy event and
has a tegional significance within the orogenic
belt.
The late Fy thrusts developed soon alter
the pewk metamorphic temperatuses had been
treuched and movements continucd during
waning Iumperatures. Some strongly schistese
retrograde rocks have formed within the thrust
zones, including the interesting Williamstown
danvourile-kyanite-corundum schists, Dis-
Placements on some of these fhiwsts were
large, and in the case of the Murray Vale
thrust, blocks with considerably different struc-
tural and metamorphic character have been
brouphe into conjunction.
Thrust faults (strike faults, overthrusts) have
beeo described from various parts of the Met.
Lolty Ranges, but us yet few have been ade-
Yitately mapped. Howchin (1906) recognised
an vyerthrust helt in the foothills of the Mt.
Lofiy Ranges near Adelaide and envisaged an
cast lo west movement. Sprigg (1946)
desenbed this belt in greater detail and con-
cluded that the amount of over-tiding was
small. Thrusts und strike faults are also well
established on the western side of the
Houghton Inlicc (Benson 1909, p. 105; Spry
(951; Campana ef a/, 1955). Freytag ( 1957)"
recoynised and mapped several strike faults
north of Williamstown and his Enterprise
Fault may eventually prove to be a continuar
ton ef the Murray Vale Thrust of the present
survey,
A portion al the Wilhamstown-Meudows
Fault has been closely mapped in the present
survey. I'he outcrop trace o/ the fault surface
provides evidence of a stcep easterly dip and
the fault may be classed as a steep reverse
type (see also Thomson, jn Parkin 1969.
p. 108), The actual displacement directions and
the amounts of displacement are unknown,
but a dip-slip component of 3+5,000 metres
dues nop seem incongruous with the observed
metamorphic grade differences across. this por-
tion of the fault surface. Recrystallisation of
dolomitic marbles in the lower grade block
aujacenr to the Cuult surface suggests heat
derived from fault movement or from a still
warm uplifted higher grade eastern block,
Erratic schistose blocks carried within the
fault zone carry a metamorphic mineral assem-
blagé suggestive of lower amphibolite facies
K. J. MILLS
conditions. It is sugested that movement! on
this fault occurred during waning metamor-
phism in lower Palaeozoic time, There is no
evidence of any more recent rejuvenation of
movement on this fault and the present fault
scarp is 4 geomorphological erosional feature,
Further investigation will be necessary to
establish whether the economically important
Ukaparinga Fault is a branch of the Williams-
town-Meadows Fault or & steep thrust of the
post-F. type,
(4) Willianistown Window
A great deal of confusion seems to have
arisen in the previous literature regarding the
interpretation and significance of the coarse
grained schists ond gneisses in the Williains-
town-Warren Reservoir region. These rocks
were initially regarded us basement rocks by
Howchin (1906, 1926), bur Later workers (e.g.
Alderman 1942, Campana 1953, Campana
ev al, 1955) considered them to be of meta-
somatic origin. Campana (1953) mapped them
as the “Aluminous metasomati¢ zone of South
Warren Reservoir’ on the Gawler 1-mile
Geological Sheet.
The present study has establixhed that these
Bieisses are in part basement gneisses of the
Warren Inlier and in patt metamorphic
equivalents of the ower portion of the Ade-
laidcan succession. The present author inclines
to the view that the schists ancé gneisses may
be the result of an isochemical reconstitution
of initially alminous pelites of mot unusual
composition under high grade metamorphic
conditions within the — sillimanite-muscovile
zone of the amphibolite facies. Vheir striking
texture pnd coarse grain-size is a result of
syntectonic crystallisation mear the peak tem-
perature of metamorphism within a zone of
lacally intense [5 deformation. Adjacent meta-
sandstones, although showing remarkably well-
preserved cross-bedding, and cyen some clastic
grain textures, huve, in fact. been subjected tes
similar metamorphic conditions. Later Fs up-
thrusting of the Warren Inlier und ifs sur-
rounding mantle of high grade Adclaidean
metasediments, principally on the Murray Vale
Thrust, and later updoming in the Fj phase
and subsequent erosion has exposed a unique
kind of tectonic windew within which we can
examine the metamorphic and structural re-
constitution of the pre-Adelaidean hasement
" Freytag, FB ('947)—The Vietoria Creek Marble, with observations of the geology of an area north-
east Of Williamstown, South Australia, Unt. Adelaide. Honours B.Sc, 'T
hesis.
STRUCTURAL GEOLOGY OF WARREN NATIONAL PARK
and the lower levels of the Adelaidean suc-
cession in the core of the orogenic belt,
fe is proposed to call this structure the
Williamstown Window. Rock units within the
window comprise the basement of the Warren
Inlier, the basal metasandstones in the Wil-
liamstown region, and the overlying coarse-
grained Adefaidean schists and onersses
belonging to Campana’s "Aluminous meta-
samatic zene of South Warren Reservoir”,
(5) Economie Considerations
li seems apposite to present here a few
comments on the structural conirul of same
af the economic mineral concentrations within
the area surveyed.
The Ukaparinga (Unapannga) Copper
Mine, Jocated in the north-west corner of the
mapped area, was worked in 1350 as the
Wheal Fricndship Mine (Brown 1908, p, 137:
Cornelius 1940) although apparently littl ore
was won. Thete hus been some renewed in-
terest in this deposit as 4 low-grade copper
prospect in recent years (Blissett 1965). This
deposi is located within the Ukaparingu Faull
zone which has been traced south of the mime
property, From Blissett’s description of the
mineralized zone within the mine the fault
zone would appear to be dipping 65-70° east,
and this fault is a steep reverse type with a
very large component of displacenvent, A
point uf some interest is that the thick tremo-
litic marble bed mapped in the South Para
yorge, suuth of the mine, can be predicted to
occur within the footwall block bencath the
mine, and this might portend a more richly
mineralized zone at depth. Tt is not yet knowa
whether the Ukaparinga Fault is a branch of
the Williamstown-Meadows Fault or is a late
F.. thrust fault. Several other copper prospects
are located within sinnlar fault zones north of
Williamstown.
The short-lived Gumeracha and Muunt
Crawford Goldfields, located uround Watts
Gully in the Mount Crawlord State Forest,
were discovered in 1884 and developed as an
alluvial field. Numerous shafts were sunk in
an attempt to locate the source of the gold,
hut the gold was apparently confined ta quartz
leaders of small size which proved utpayable
(Brown & Woodward 1886), The known gold
distribution in this field is confined to the hinge
region of a large F, antiform, the Gumeracha
Goldficlds Antiform, and it scems likely that
the auriferous quarts leaders were loculised
wilting (ractires associaled with this brittle
Jute metamorphic fold structure. This struc-
J|3
tural control may prove to be of some interest
in the interpretation ol the distribiillon of
other goldfields im the Mt, Lo'ty Ranges,
The Williamstown clay deposits have been
worked in the vicinity of the Warren Reservoir
since early this century, Several older disused
mines are located south of the reservoir within
the mapped area, especially within north-south
zones of highly altered rocks running immedi-
alely east of the Wirrianda homestead and east
of the road immediately south of the old road
bridge, buf the main quarry currently operated
is that in section 950, Hundred of Barossa,
just outside the north-cast edge of the mapped
area. Many reports have been wrillen On these
deposits but hitherto no important structural
control has been recognised. Jack (L926) sug-
gested that the kasolinisation and hydromica
development was preater than would be
expected by weathering alone and suggested
some Form of introduction. Cornelius (1932)
indicated that the deposit in section 950
dipped steeply east and had a southerly pitch.
Alderman (1942, 1950) concluded that the
cluy deposits had largely originated through
the hydration of an original body of massive
decussately textured sillimamite rock and
erected an elaborate metasomatic scheme to
explain the various phases present and the
sudden changes from low grade to high grade
rocks. Alderman (1942, p. 8) states that “there
dovs not scum to be any major structural
feature separating the aluminous rocks Irom
the low grade rocks”, Later workers (Gaskin
& Sampson 1951, p. 60; Betheras 1953;
Cochrane 1954, p. 54: Cochrane 1955, ». $5)
generally accepted Alderman’s views, and
although there are references fo various
damourite shears and small faults, these are
cunsidered to be of minor importance. and
perhaps cue to volume changes during metu-
somatism, The clay deposits appear to be
irregular in distribution and this was related
fo the fortuitousness of metasomatism.
The clay deposits north of the reservoir
were not studicd by the author, but pre-
liminary observations have tndicaled that the
Murray Vale Thrust passes through the centre
of the large quarry on section 950 and
separates the highly altered sillimanite-kyaqite
bearing rocks to the west front the low-grade
“clay-states" to the east. It the vicinity of
the quarry the Murray Vale Thrust is ap-
parently involved in Several tight F, folds,
Tn terms of the structural history deter-
mined for the schists occurring south of the
js K, J. MILLS
reservoir, the following series of events is
envisaged us controjling (actors in the forma-
tion of the clay deposits, The original mussive
sillimanite-rutile rocks are thought to have
crystallised within the sillimanile-nmiuscovite
zone schists and gneisses at the peak of metu-
morphism under static conditions during the
interkinematic period between the F, and F,
deformation events, The origin of the silli-
manite remains obscure, but presumably same
form of melasomatic transfer was involved.
During the Fy, event, whilst the country rocks
were being intensely deformed and reconsti-
tuted. a number of damourite-kyaniles. corun-
dum shear zones developed within the tectonic-
ally resistant sillimanite pods and late in the
F, event several strong thrust zones appeared,
and the subsequent passage of hot aqueous
solutions within the thrust zones altered the
sillimanite deposits to clay. It is considered
that the sillimanite and damonrite-kyanite
deposits may be structurally controlled within
early incipient zones of [F. Uhrusting, but this
will need further investigation. The subsequent
development of clay within some of the silli-
manite bodies may have required the presence
of open channelways along zones of tuter
movernent and could be enhanced by the deve-
lopment of strong Fy folds in the vicinity of
section 950,
(6} Sanpnary
The following list of events summarises the
proposed geological histary for the Warren
Reservoir region:—
{a) Deposition of pre-Adelaidean sediments
of the Warren Inlier. These sediments were
largely pelitic and homogeneous with some
quartzofelspathic units and minor mars.
(b) Deformation of the pre-Adclaidcan sedi-
ments of the Warren Unlier. Several phases of
folding, as yet unravelled, accompanied this
deformation, with metamorphism (o upper
greenschist or lower amphibolite facies at
most, There gre no intrusive tocks, apart from
pegmatices. no volcanics, no rocks of
“Houghton diorite” type, and no evidence for
a phase of pre-Adelaidean retrogression and
phylonitisation,
(c) Krosion to 4a peneplain surface,
(d) Deposition of the Adelaidean sequence,
consisting of a basal cross-beddled felspathic
sandstone followed by a pelitic sequence grad-
ing upwards into more calcareous or dolo-
mitic sequences.
(c) Deformation and metamorphism of the
Adelaidean succession in the lower Ordovician
Delamerian orogeny. Three compressive events
accompanied this orageny and metamorphism
locally reached the sillimasite-muscovile zone
of the amphibolite facies within and adjacent
to the Warreb Inlier A locally important
phase of thrusting from the east accompanied
and followed the second compressive event
under conditions of waning metamorphic tem-
peratures. The third compressive event, ol
more brittle character, occurred during low
metamorphic lemperutures and played an im-
portant role in the development of various
mineral deposits,
(1) Formation of the Wilhamstown-
Meadows Fault as a major plane of discon-
linuily extending along most of the length of
the central Mount Lofty Ranges. ‘This fault ts
responsible for bringing the higher grade meta-
morphic zones of the eastern Mount Lolly
Ranges into conjunction with the lower grade
zones of the western Muunt Lofty Ranges.
Displacement on this fault occurred in the
early Palacozoic and there is no evidence of
subsequent rejuvenation of movements.
(zg) Extensive denudation and peneplanation
and development of a marked fault scarp ero-
sional feature along the Williamstown.
Meadows Fault in the Williamstown region
through the differential erosion of (he lower
grade phyllites by the South Para River.
Acknowledgements
The author is grateful for the co-aperation
of the various public autboritics and private
individuals who have permitied access to pro-
perty fo make this study possible, The project
Was completed in the Department of Geology
and Geophysics, University of Sydney. Mr. R-
Sealy kindly axsisled with photographic work
associated with the construction of the figures.
Mrs. J, Neilsen as especially thanked for her
patient redrafting of figure t. Mr, Glenn
Kulson yssistel with the lettering on figure 2
and the map legends, Mrs. R. McKenzic
typed the final manuscript. The venerous sup-
port of the University of Sydney in enabling
the completion of this work is gratefully
acknowledged.
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OBITUARY: THEODORE GEORGE BENTLEY OSBORN,
D.SC., M.A., F.L.S.
Summary
THEODORE GEORGE BENTLEY OSBORN
D.SC., M.A., F.L.S,
OBITUARY
THEODORE GEORGE BENTLEY OSBORN, D.Sc., M.A,, F.L.S.
2.x.1887 -3.vi.1973
Theodore George Bentley Osborn was born
in England and educated at Burnley Grammar
School and the University of Manchester. He
spent just over half of a long and energetic life
in Avstralia, occupying Chairs of Botany at
Adelaide and Sydney Universities and later the
Sherardian Chair at Oxford University, On
retirement he chose to return to Adelaide.
Osborn's research during the period 190#-
1912 was on the fungi, when he was u lecturer
in Economic Botany in the University of Man-
chester, In 1911 he was awarded the M.Se.
degree, and in 1920 the D.Sc. degree, from the
same University.
In 1912, Oshoen was invited to become Pro-
fessor of Botany, Vegetable Pathology and
Parasitology in the University of Adelaide,
eombined with the office of Consulting Botanist
to the Government of South Australia, and his
publications over the next nine years strongly
reflect the nature of this appointment,
Shortly before coming tu Australia, he mar-
ned Edith Muy Kershaw, M,Sc-., Assistant Lec-
turer in Botany in the University of Manches-
ter. They had three sons, Peter, Andrew, and
Richard, all of whom served in World War IT;
Andrew, an R.A.F, pilot, was killed in action
in 1942,
When he established the Department of
Botany in Adelaide in 1912, Osborn was only
25 years old and youthful in appearance, He
was fond of the story of how he went into the
examination hall, which was being supervised
by a middle-aged woman who had not met him,
and asked for the Botany I paper. She brought
him the paper and when he said “and while 'm
here [It have a look at the Agricultural Botany
paper too" she replied, “You'll do nothing of
the sort, young man, you'll just sit down there
and get on with your Botany I.”
Osborn developed the first degree courses in
Botany at Adelaide University, though Botany
courses had been given earlier under Professor
Ralph Tate. He took most of the teaching until
1916 when a demonstrator was appointed, fol-
lowed in 1922 by a lecturer in Plant Pathology.
The latter appointment, and the establishment
of the Waite Agricultural Research Institute in
1925, relieved Oshorn from much of his time-
consuming advisory work. However, in 1926
he was asked by the executive of the recently
formed Commonwealth Council for Scientific
and Industrial Research to review botanical
work in progress in Ausiralia, and further to
report on plant problems in Australia. In. 1927
he was appointed adviser to the Council and
was offered the dircctorship of their Division
of Plant Industry. However, at about this time
he applicd for the Chair of Botany at the Uni-
versity of Sydney, lo which he was appointed
in 1928.
During the early years of his residence in
Adelaide, Osborn deVeloped interests in plant
ecology, studying areas near Adelaide as well
as distant ones, stich as Franklin Island and the
Pearson Islands, off the west coast of Eyre
Peninsula. During a visit by Professor R. S.
Adamson (University of Cape Town), joint
studies were made of the ecalogy of the Ooldea
district and of Eucalypries Forests of the Mount
Lofty Ranges. ‘These. studies: were published in
the Transactions of the Royal Society of South
Australia. Osborn also published on two primi-
tive lycopods, (Isoetes and Phylloglosswn),
found for the first time in South Australia.
These ecological studies led to Osborn's great
interest in the vast arid region of South Aus-
tralia. He made numerous visits to the saltbush
areas and developed cordial relationships with
pastoralists. This culminated in the generous
gift of Messrs. Hamilton, Wilcox Ltd., in 1925,
of the Koonamore Vegetation Reserve, which
is now the oldest biological station, with con-
tinuous records, in Austrolia, The reset've,
‘Trans, HM, Soc. S. Aust, Vol 97, Part 4, 30 November, 1973
318
located 385 km NNE of Adelaide, was re-
named the “T, G, B. Osborn Vegetation
Reserve at Koonamare” i) |972. The first
important work on grazing and regeneration of
natural Vegetation its chese arid regians resulted
from the collaborative work of Osborn and his
two colleagues. J. G. Wood and T. B- Paltridge.
The reserve and nearby areas are still used
extensively by stall and students of the Ade-
laide Botany Department.
in 1928 Osborn became Professor of Botany
at the University of Sydney and he set about
rvorganising the undergraduate courses there.
He took the first year lectures and those in
plant physiology, as well as in ecology. At first
he continued to clear wp his ared zone research,
relaining direction of the Koonamure work
uotil 193), but increasingly he became in-
terested in the vegetation of coastal New South
Wales, In [930 he gave the Livingstone Lee-
Iures enuiled “Plaot Life in the Sydney Dis-
trict", with an ecological upproach. He was
interested in the xerophytic propertics of the
plants characteristic of this relatively high Tain-
fall area. In 1932, his Presidential Address to
the Linnean Socicty of New South Wales. was
on “The Plant i Relation to Water’, with spe-
cial reference to the properties of xerophytes
in being able to withsiand drought. Later, O3-
horns work itt ecology stimulated the interest
of his students in descriptive ecological
accounts of other parts of New South Wales,
including one on the ecology of the Myall
Lakes vegetation, jointly with R. N, Roherlson.
During his time in Australia, Osborn was an
aclive member of 4cientific societies. He was
4 stroug supporter of the Royal Society of
South Australia, contrihuting several papers
uml berng a Fellow since 1913 and Honorary
Fellow since 1955. He was a Council Member
1915-20, 1922-24, Vice-President 1924-25,
1926-27, anu President 1925-26. He played an
active part in the establishment of Flinders
Chase on Kungaroo Island, being a member of
the board from 1919 to 1927, and Honorury
Secretary and Treasurer 1921-25. Oshom was
alsa a member of the Linnean Society of New
South Wales from 1928 and President in 1932,
and President of Section M (Botany) of
A.N.Z.A.AS. in 1928, and a Vice-President
from 1924-25, The Royal Society of New
South Wales awarded him the Clarke Medal in
1958.
Osborn became Sherardian Professor of
Holany aid a Fellow of Magdalen College, Ox-
ford, in 1937, The Oxford Department was
very poorly housed in cramped surroundings in
an antiquated building in the Oxford Botanic
Giarden. This Bolanic Garden, founded in
1621, had been somewhat neglected. but Os-
born stinvulated collaborative work with the
Garden, which continued strongly after the
Department moved into the new laboratories
planned by him and consteucted shortly before
his retirement in 1953.
Oshorn's achievements ut Oxford have been
summarised in his obituary in TRe Tintes for
6 June 1973~—-There can be nu doubt that
under him the study of botany m Oxford
achieved an eminence which it had not before
reached even under more famous Sherarlian
Professors. Not only was the production of
original work in botany greater in quantily and
quality than ever before but of the undergra-
duates, demonstrators and research men pre-
sent in Oxford during his tenure of the chair,
eleven of more are or have been Professors
and heads of departments, and several mure
Readers in biological departments of Univer-
sitics in Britain and clsew here.”
In 1953, Osborn retired from the Chair at
Oxford sad returned to Australia, After a
period of residence in Adelaide, in 1957 the
Osborns moved to Melbourne where Mrs. Os-
born dicd a year later. Following this, Osbora
returned to Adelaide to live. During 1959 he
was Acting Master of St, Mark's College, of
which he had been one of the founders in the
carly 1920's, In 1962, when the Botany Depart-
ment celebrated its 50th anniversary, he was
appointed Professor Emeritus of the University
of Adelaide, having been given a similar title
at Oxford University on his retirement in 1953,
Osborn left Oxford well known, among other
things, for his course on Gyminosperms, ane in
Adelaide he was invited by Penfessor J. G.
Wood to give 4 course oo this group of plants
to third year students. This he contrnusd to do
for five or six years, and in 1960 published his
last paper, on the embryology and life history
of Podevarpus faleatus. Osborn had long in-
tended writing a book on the Gymnosperms,
but unfortunately this never came to fruition
Osborn made his home at St. Mark's College
during 1959, where he was associated with the
very capable seerctary Marjoric Sabine, whom
he married in England in 1960. Both Professor
and Mrs, Osborn took an active part in the life
of the Botany Department from then on, and
Osborn attended the weckly seminars until
shortly before his death,
Oshorn was a good lecturer and teacher who
attyacted students of quality, His influence was
passed on toa several of the present leaders in
Australian plant ecology, and among his many
Australian students were: J. G, Wood (for-
merly Professor of Botany, Usmversity of Ade
lade}, Ci. Samuel (formerly Deputy Chiel
Szientific Officer of the Agricultural Research
Council, WK), G. J. Rodger (formerly
Direztor-General of Forestry, Canberra), B, H.
Bednall (formerly Conservator of Forests,
South Australia), T. B. Paltridge (formerly
Chief Scientific Liaison Officer, C.S.1.R,0,),
N. A. Burges (formerly Professor of Botany,
University of Sydney, and Vice-Chancellor,
University of Northern Irclandi, M.R. Jacobs
(formerly Director-General of Forestry, Can-
berra). D. Martin (Officer-in-Charge, Tis-
mangn Regional Laboratory, C.S.1R.O.),
H. K. C. Mair (formerly Director and Chief
Botanist, National Herbarium, New South
Wales), BR. N. Robertson (Director, Research
School of Biotogical Sciences, Australian Na-
31y
tional University), Lilian Fraser (formerly:
Chicf Biologist, Division of Science Services,
N.S.W, Department of Agriculture), Joyee
Vickery (formerly Scoior Botantsi, N.SW,
Department of Agriculture), N.C. W. Beadle
(formerly Professor of Botany, University of
New England), Gwenda Davis (formerly Asso-
claie Professor uf Botany, University of New
England), N, H. White (formerly Professor of
Plant Pathology, University of Sydney), and
Ilnva Brewer (née Pidgeon, School of Bio-
logical Scienzes, University of Sydney).
Osborn died after a short illness on 3 June
1973. He will be long remembered in Australia
Yor the mfluence he had on Botany im Adelaide
and Sydney, and particularly for his qualities
of leadership and inspiration of younger
people, He always had the huppy knack, even
m his later years, of being able to talk with
Students, lo be interested in what they were
doing, und to cutch their interest in his ex-
periences.
—R. N. Robertson and C. M- Eardley.
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