THE SECOND CENTURY

The Royal Society of London was conceived more than 300 years ago when, following

a lecture by Christopher Wren, it is recorded that “something was offered about a designe of

founding a Colledge for the promoting of Physico-Mathematicall, Experimental] Learning”.

Charles II became interested and granted the Society its first Charter in 1662, naming it “The

Royal Society”. A second Charter, in 1663, granted the body Arms, bearing the motto

“Nullius in verba”, as an expression of the determination of the Society to resist all dogma

and to verify all information and statements by appeal to observation. At the same time, the

full title of the Society became “The Royal Society of London for Improving Natural Know-

ledge”.

Sir Mark Oliphant, K.B.E., Patron 0} the

Royal Society of South Australia

A little more than 200 years later, the Royal Society of South Australia was born, and

1976 is the centenary of its Transactions. Throughout this time the South Australian Society

has played an invaluable role in recording, in proper scientific manner, manifold observa-

tions and descriptions of the geology, the fauna and the flora of this State. For long, the

Society has emulated in its activities the search for natural knowledge which was the objec-

tive of the founders of the original Royal Society. However, for very practical reasons, such

bodies are reappraising their activities and ever questioning whether they can continue to

exist. Strangely enough, the problems arise from the very success of their past work. The

technology, the application of knowledge for practical purposes, which arises from increasing

understanding of nature, has made of science an ever growing national and international

activity. Through their control of money, governments now determine the level of scientific

enquiry and its content, while the rapidly increasing cost of publication means that they

become the arbiters of what science shall be printed.

SIR MARK OLIPHANT

There are other factors which have a pronounced influence on comparatively small

societies allempting to cater for those interested generally in science, Foremost is the growth

of specialist socicties interested only in a very marrow area of natural knowledge. It is almost

impossible today to attract a biochemist to a discussion of relativity or cosmology, or to per-

suade a nuclear physicist that the nature of the chemical bond can be an interesting subject

of study, The particle physicist cannot understand the entomologist, or the computer elec-

tronics expert the ornithologist. A broad interest in nature generally is now rure. There are

few today whose interests are as broad as those of Lord Rayleigh, who wrote learnedly of

the hending of marble mantlepieces and developed the mathematical theory of the wave-

guide, Publicution of the results of scientific enquiry becomes more and more confined to

specialist journals, or specialist sections of older journals.

Secondly, it is the national badies, like the Academies of Science of the United States and

Russia, which are gavernment agencies, or the Royal Society of London and the Australian

Academy of Science, which are not controlled directly by government. which represent

scienee internalionally for their countries, and which receive the greatest financial backing, Tt

is they Who can afford to organise conferences, national or international, they who can pro-

vide the expenses for travel by members and by visitors. They are prestigious societies whose

activities are of world-wide significance.

The Royal Society of South Australia cannot emulate these giants, However, T any

convinced that it will have @ vital part to play in the second century of its existence. The

growing interest, among scientists and the general public, in the preservation of the environ-

ment, in pollution, ecology, our national heritage of fauna and flora, natural resources, and

the beauty of landscape, encourages belief jn a return to deep interest in nature generally.

rather than in ohe aspect only, Tf, as C believe, our Royal Society is ta be revitalized to

hecome a significant social influence through scien¢e, it must espouse catlises, after full and

frank discussion to determine an agreed approach, and then speak loudly and in public of its

conclusions,

As mstances of broad issues to which the Society could contribute much, there are

preservation of the Mf Lofty Ranges and the Flinders Ranges, the salinity of the River

Murray, the deterioration of our desert areas, We should not be deterred by the existence of

CS.LR.0., of State instrumentalities, or of the Australian Academy of Science, from

choosing areas of comprehensive, interdisciplingary study, which can be exciting scientifically

and could be rewarding socially. A paper describing a new species of coleoptera from the

Lake Frome area should be published, but will not rouse much interest in a general scieutitic

audience. On the other hand, if this discovery is significant in indicating climalic change, or

ecological upset On the castern borders of the Flinders Ranges, it could become an important

contribution to wide discussion,

South Australia faces a multitude of problems, most of which are relegated to investi-

galian by governmental bodies. Qur tuiure depends erilically upon assured supplies. of water

and of energy. Water, gas, and elcetricity authorities are too busy with immediate questions

to exercise the imaginative resypanses necessary for creation of new approuches to longer

term problems. On the other hand, a Royal Saciety which stimulates constructive discussion

of such problems, in their widest context, might generate completely new ideas.

Where are We going? Thatis the first questiam we must answer in the early part of our

second hundred years.

GovERNOR oF Sourht AUSTRALIA

COMPARATIVE OSTEOLOGY OF THE PELVIC GIRDLE OF

AUSTRALIAN FROGS AND DESCRIPTION OF A NEW FOSSIL GENUS

BY M. J. TYLER*

Summary

TYLER, M. J. (1976).-Comparative osteology of the pelvic girdle of Australian frogs and

description of a new fossil genus. Trans. R. Soc. S. Aust. 100(1), 3-14, 28 February, 1976.

The osteological characteristics of the pelvic girdle of twenty-five extant genera of Australian frogs

of the families Hylidae, Leptodactylidae, Microhylidae and Ranidae are defined. The new Tertiary

fossil genus and species Australobatrachus ilius are described from the Etadunna Formation.

The fossil exhibits a unique lateral ilial groove and is referred tentatively to the family Hylidae.

COMPARATIVE OSTEOLOGY OF THE PELVIC GIRDLE OF AUSTRALIAN

PROGS AND DESCRIPTION OF A NEW FOSSIL GENUS

by M. J. TyLer*

Summary

‘Tyter, M. JI. (1976)—Comparative ostealogy af the pelvic girdle of Australian frogs and

description of a few fossil genus. Trans. R. Sac. 8, Aust, 100(1), 3-14, 28 February,

1976.

The osteological characteristics of the pelvic girdle of twenty-five extant genera of Aus-

tralian Trogs of the families Hylidae, Leptodactylidae, Microhylidac and Ranidge are defined.

The new Tertiary fossil genus and species Australobutrachus ilius are described from the

Fiadunna Formation, ‘Ihe fossil exhibits a unique lateral ilial groove and is referred tentatively

to the family Hylidae,

Introduction

Of all bones of the anuran skeleton, the

ilium has been shown to vary considerably and

consistently between families, genera and even

Species, Ilia are commonly well preserved

amongst disarticulated skeletal material, and

their features are sufficiently diagnostic to per-

mil identification; for this reason ilia have pro-

vided the basis for the recognition of genera

and erection of new fossil species: (Lynch 1963;

Chantell 1964; Holman 1965),

Dala on extant species so essential for identi-

fication and general comparative purposes are

frequently limited, and in the case of Australian

frogs, data are particularly deficient. Lynch

(1971) provides the only comparative contri-

bution. Confining his interest to leptodactylids,

Lynch described the ilia of representatives of

nine genera. Since then a fossil ilium of a pre-

viously undescribed genus bas been reported

from the Australian Tertiary by Tyler (1974).

uml it has proved necessary to examine und

describe representative ilia of all of the known

living genera in order to describe the new genus

and species,

Material and methods

The dry specimens of the modern species

studied were dissected from representatives of

4 families, 25 genera and 60 species, including

ull geneta known in Australia, This material is

in the author's collection. The fossil described

herein is in the Palaeontology collection of the

South Australian Museum.

With only minor variation, the descriptive

terminology used follows Lynch (1971), and

the features recognised are shown in Figure 1,

Morphometric data were obtained with dial

callipers or an eyepiece micrometer. The length

of the animals from snout to verit Was measured

before dissection. Subsequently, the distances

between the tip of the dorsal acetabular expan-

sion atid the end of the ilial shaft and the span

between the anterior margin of the dorsal pro-

minence and the ventral acetabular expansion

were measured, The bone measurements were

examined to establish relationships between ilial

size Or proportions and the size of the donor

frog.

Features of the Anuran pelvis

(A) Puhis

The pubis is customarily a small, roughly tri-

angular, cartilaginous wedge of tissue separat-

ing and underlying the ventral borders of the

ischium and ilium (Fig. 1). In particularly

large specics. (rarely in small ones), where

there js a more intimate degree of fusion of the

pelvic components, the pubis is often calcified

or ossified.

(B) lyehinm

The ischium is .a bony or cartilaginous disc

anteriorly fusing with the ilium to provide the

posterior half of the acetabulum, 4nd pos-

* South Australian Museum, Nocth Terrace, Adelaide, S. Aust. SOD0. (Present address Department of

Zoology. University of Adelaide, Adelaide, ‘S. Aust, 30011),

4 M. 7. TYLER

Nial

shaft Oprom. Dacetexp

D protub, ae

\ p »

pre-acet, Zone

Vacet.exp, Ischium

Pubis

Acet. fossa

Fig, 1, Left lateral aspect of anuran pelvic girdle.

Abbreviations: Acct, fossa-—aceiabular

fossa; 0. acet. exp—dorsal acetabular

expansion; D. prom.—dorsal prominence:

D. protub,-—dorsal protuberance; pre-iacct.

zone—pre-acetubulur zone; V. acel. exp.—

ventral acetabular expansion.

teriorly forming a plate occupied by numerous

mus¢les communicating with the femur,

The shape of the ischium varies considerably

at a specific level and is associated with dif-

ferences in habits. For example, the develop-

ment of a particularly large plate posterior to

the acetabular fossa ts found only in rotund

apecies with exceptionally short and muscular

hind limbs. Similarly, the development of a

high, dorsally projecting extension is charac-

teristic of large but ugile frogs with powerful

hindlimbs. Viewed with the femur as the

reference point it is clear that a major shift in

the inuscle mass can only be brought about by

voncurtent changes in the development of the

ischium.

The ilium is the, largest pelvic bone, and con-

sists of an elongate shaft terminating in an axe~

head shaped body. It is a paired stricture

articulating anteriorly with the ventral surface

of the sacra] diapophysis and posteriorly abut-

ting the ischium and pubis. Various areas of the

tlium ate recognisable as distinct components -

(a) MMial shaft: The ilial shaft is an elongate

and usually slightly curved structure varying

from «a cylindrical section through vertically

oval to more elaborate forms in which there

are grooves on the lateral or medial surfaces.

(b) Dorsal crest: A smooth bordered crest, ris-

ing high as a thin blade, occurs in numerous

species of frogs, but is rare amongst Australian

forms. It usually arises from the dorsolateral

surface of the shaft, and reaches its maximum

height within the anterior one-third of the shaft

(Fig. 4L),

is a diflerentiated area rising on the superior

margin of the shaft in a position above or

slightly anterior to the acetabulum. [t is not

present in all species and Js scarcely detectablc

in many others (¢.z. in those exhibiting a dor-

sal crest)_

(d) Dorsal protuberance; From the body of

the dorsal prominence the dorsal protuberance

arises as an elongate, rounded or pointed knab.

To it uttaches the Muscutus gluteus maxirnus.

(e) Dorsal agetabular expansion: The body of

the iliim extends dorsally into. a triangular por-

tion of bone superior and posterior to .the

acetabulum. This projection is termed the dor-

sal acetabular expansion. It may rise steeply

atid so meet the ischium in a vertical plane,

whilst the anterior face can be at an acute or

obtuse angle to the ilial shaft,

(f) Acerabulum: Variation in the acetabulum

consists of differences in size (relative to adja-

cent structures), in position in relation to the

ilial shaft, and in the width and extent of

development of the acetabular rim,

(g) Ventral acetabular expansion: The superior

segment of the ventral acetabular expansion

has heen termed the preacetabular zone by

Lynch (1971), The considerable variation in

this portion of the pelvis is difficult to express,

because of the instability of potential reference

points such as the ucetabulum, Nevertheless,

the basic shapes range from the form of a

straight line extending ventrally and posteriorly

(Fig. 24, A), a gradual concavity (Fig. 2B},

or a concavity of the preacetabular zone and

a convexity bencati (Figure 2C).

Se =

A YY B ( iy

= a

C ( D “Sy .

Fig. 2. Variation in the shape of the ventral

Acetabular expansion viewed from left

aspect. See text for explanation,

PELVIC GIRDLE OF AUSTRALIAN FROGS

Fig. 3. Pelvis or isolated ilium of hylid and leptodactytid frogs:

A, Litoria caerulea, x 2; B. L. lesueuri, x2; C. L, eucnemis, x 5; D. Nyetimystes zwiefeli, x 2:

E. Adelotus brevis, x 5; F. Assa durlingtoni, x 5; G. Crinia georgiana, x 5+ HH. Cyclorana nevae-

hallandiae, x 2; I. Geocrinia laevis, x 5; J, Glauertia orientalis, x 5; K, Heleioparus albopunctatus,

x 2; L. Kyarranus sp., x 5; M. Lechriodus fletcheri, x 5; N. Limnodynastes peroni, x 5; O. Mixo-

phyes fasciolatus, x 2.

POST-MORTEM CHANGES

Disintegration of the pelvic girdle into its

component bones has been observed in species

in which cartilage is most extensive. Species

that are heavily ossified remain intact.

During the process of dehydration, the

acetabular fossa may become distorted and,

in the material available, such distortion takes

the form of compression in « dorsoventral

direction. The dorsal crest of the ilium is par-

G M, J. TYLER

TABLE 1

Characteristics af ihe Uiant tn species a} Litoria

Dorsal pro-

minence/ Acetabular Position

Profile of yentral anterior rim rim ventral of medial

acetubular of ilial shaft rim on

Species expansion Acetabulim margin Pubis Tlivm

adelaidensis concaye level above curt, ant, t

qneiana* concayo-conyex posterior level cart. absent

dured concave 4 antenor above curt. ant. 7

hicoler concave 4 anterior above cart, Tabatnt

hoeraolangertsis concave-canvex 4 anterior level bony central 1

brevipalmaia concave level abave curt, ant, Ly

caerulea concave level level curt. ubsent

citropa concive level above cart, ant.

darsalis® concavo-convex 4 anterior level curt. aot. ¢

eucnemis concavo-convex level above bony ant. 4

ewingi concave level above curt Post. +

gracileita concavo-conyex 3 anterior above cart. absent

infrafrenata concaye + anterior above cart. anh. 3

lesiteuri concavo-convex + antetior level bony central 43

microbelos concwo-convex level above Gurl. ant, 4

nannotis concaye-conyex level above bony absent

nasula concavo-canyex 4 anterior above carl. ant. 14

nizrofrenata concayo-conyvex \ amterior above bony ant, $

rubella concave 4 anterior above vart. absent

rothi concave + qnlerjor above curt. absent

thesaurensis* conceave-convex leyel above bony ant, 3

ant—anterior; cart—aartilaginous.

ticularly subject to post-mortem distortion,

commonly bending medially from a perfectly

vertical orientation to form a quadrant, Even

more conspicuous is the distortion amongst

material recovered fram owl pellets where

there has been an induced medial curvature of

the ilial shaft in. several specimens.

Account of modern genera and species

Family HYLIDAE

LITORIA Tschudi

FIG. 34-C

Species examined: L, adelaidensiy (Gray). L.

angiana (Boulenger), L. aurea (Lesson), LZ,

bicolor (Gray), L. boaroalongensis (Moore),

I... hrevipalimata Tyler, Martin & Watson, E.

caerulea (White). L. citrapa (Tschudi). L. dor-

salixs Macleay, L. eucnemis (Lonnberg), L,

ewingi Dumeril & Bibron, ZL. gracilenta

(Peters), L. infrafrenara (Gunther), D. lesneuri

Dumieril & Bibron. L, micrebelos (Cogger). L,

nannotis: (Andersson), £, naswia (Gray), L.

nigrofrenata (Gunther), 1. rubella (Gray), L,

rothi (de Vis), L. thesaurensis (Poters).

Variation in this morphologically and ecola-

gically diverse genus renders wa generic defini-

tion a difficult proposition. For this reason

comparative data arc included in Table 1, and

only the following generalisations ure possible:

The pubis is cartilaginous or ossified and the

ischium is ossified.

The ilial shaft lacks a dorsal crest but in-

variably bears a narrow tim on at least a por-

tion of the medial surface. In L. aurea there

is also a lateral groove exhibiting a distinct

ontogenetic trend in hecoming progressively

less conspicuous, The acetabular Jossa tends to

be rather large. The ventral acetabular expan-

sion is of a varicty of forms, from a narrow,

concave profile to concave-convex,

The dorsal protuberance and dorsal

prominence are usually well differentiated but

are not raised high above the level of the ilial

shaft.

NYCTIMYSTES Stejneger

FIG. 3D

Species examined: N, tympanocryptis (Ander-

sson and N. zweifel? Tyler of New Guinea.

PELVIC GIRDLE OF AUSTRALIAN FROGS 7

TABLE 2

Generic features of Tlia

Dorsal prer-

minence/

lial shaft Ital shaft Dorsal anterior rim

cresh rim protuberance of acétabulum

Adelotus. absent present prominent anterigr

Assa absent absent inconspicuous level

Caphixalus absent absent inconspicuous anterior

Crinia absent present inconspicuous anteriar

Cyclorana absent present or absent incanspicuous anterior

Geocrinia absent absent absent level

Glauertia absent absenol inconspicuous anterior

Heleioports absent absent prominent anterior

Kyarranus absent present prominent anterior

Lechriodus present absent moderate posterjor

Limnodynastes present or absent present or absent prominent ant. or Jevel

Litoria absent present or absent moderate usually anterior

Mixophyes present absent inconspicuous anterior

Myobulrachus absent absent absent posterior

Neobatrachus absent absent prominent anterior

Notaden absent absent prominent posterior

Nyctinystes absent absent moderate level

Philoria absent. present inconspicuous anterior

Pseudophryne absent absent moderate level

Rana present absent inconspicuous posterior

Ranidella absent absent moderate anterior

Rheobatrachus absent absent prominent posterior

Sphenophryne absent absent iNconspicuous posterior

Taudactylus absent absent prominent anterior

Uperoleia absent absent prominent anterior

—___—_—_—_——————OOOOOO Cc oO — — — —

The ilium and ischium are ossified in both

species. The pubis is ossified in zweifeli und

Cartilaginous in tympanocryptis.

The ilial shaft is long, curved distally and

very slightly compressed mediolaterally. The

ventral acetabular expansion is gently rounded

in a single, uninterrupted concave arc. The

acetabular fossa is prominent, with its. upper

margin level with the centre of the ilial shaft,

The dorsal prominence and dorsal protuberance

are small and project laterally rather than

superiorly. The anterior margin of the dorsal

protuberance is on a level with the anierior

margin of the acetabular rim, The dorsal acc-

tabular expansion is only slightly raised.

Family LEPTODACTYLIDAE

ADELOTUS Ogilby

FIG, 3£

Species examined: A. brevis (Gunther).

The ischium is bony .and the pubis is entirely

cartilaginous.

The ilial shaft is distinctly curved and bears

a narrow indentation on the medial surface.

This indentation is deepest in the midsection of

the shaft. The acetabulum has a narrow

peripheral rim which superiorly is on a. level

witb or is slightly superior to the ventral surface

of the ilial shaft. The ventral acetabular expan-

sion is only slightly developed, and the preace-

tabular zone is extremely narrow. The dorsal

acetabular expansion is elongate and raised

moderately. The dorsal prominence is poorly

defined. The dorsal protuberance is extremely

large, inclined posteroventrally and is almost

entirely anterior 10 the anterior rim of the

acetabular fossa.

ASSA Tyler

FIG, 3F

Species examined: A. darlingtoni (Loveridge).

The pubis and ischium are bony except for

the portion associated with the posterior half of

the acetabular fossa.

3 M. J. TYLER

The ilial shaft is slightly curved, lacks ridpey

and indentations and is circular in cross section,

‘The acetabulum has an exceptionally Well

developed penpheral rim which superiorly is

very slightly ahove the ventral margio of ihe

ilial shalt, Lhe Ventral acetabular expansion ts

slightly develaped inte a narrow preacetabular

zone. The dorsal acetabular expansion is Very

poorly developed. The dorsal prominence is

only slightly defined. The dorsal protuberance

is small bul promlitent, its anterior margin on

o level with the uuterior margin of the scctabu-

lar rim.

CRINIA Tschudi

FIG. 3G

Species examined; C, georgiana Tschudi.

The pubis is cartilaginous and the ischium ts

oasified,

The ilial shaft is curved, flattened medio-

laterally over the posterior half and dorsoven-

trally over the anterior half. There is no dorsal

ilisl crest but there is a very slight longitudinal

medial indentation. The acetabulum is large,

has a fairly broad peripheral mim which

superiorly is very slightly above the level of the

ventral margin of the ilial shaft.. The ventral

acetabular expansion is only slightly develaped;

the subacetabular zone does not protricde an-

teriorly. The dursal acetabular expansion 13

poorly developed. The dorsal prominence is

low, but guile distinguishable from the ilial

shaft. The dorsal protuberance is just

detectable. Slightly less than one-half of the

dorsal protuberance ty anterior to the anterior

Tim of the acetabulum.

CYCLORANA Stcindachner

FIG, 3

Species examined: C. australis (Gray), C. dalili

(Boulenger}, C, novaehollandiae Steindachner

and C, platyveephalus. (Gunther).

‘The ischium is bony and fused to the thum,

whereus the pubis can be extensively ossificd

and similarly fused (C. dahli and C. navae-

hollandiae), or completely cartilaginous (©.

australis and C. platycephalus),

The ilial shaft is slightly curved and cither

hears a satrow dorsal rim, rendered con-

spicuous by a longitudinal indentation on the

medial sutface of the shaft (C. australis, C,

dalilf and C. wevaehollandiae), or elsc Jacks a

dursal rim (C. platyceplidus). The acetabulum

has a narrow peripheral rim which superiorly as

above the level of the ventral surface of the

iial shatt, The ventral acetabulac expansion ts

slightly developed and the preacetabular zone

ig narrow. The dorsal acetabular expansion is

prominent and conspicuous. The dorsal pre-

minence is distinguishable from the Wial shuti

but only slightly raised. The dorsal pro-

tuberance is inclined yentrolatersily, extends

far from the prominence, and is approximately

one-half anterior to the anterior rim ob the

acetabular Fossa.

GEOCRINIA Blake

FIG. 3/

Species examined: G, laevis (Gunther),

The pubis is cartilaginous und the ischium ts

ussified.

‘The ilial shaft is short. slightly curved and

fattened laterally in cross-section, The acetabu-

lum is moderate with a narow peripheral rim,

The superior margin of the acetabulum hes

slightly above the level of the ventral margin of

the ilal shaft. The ventral acctabular expansion

is only slightly dilated. The dorsal acetabular

expansion is not pronounced. The dorsal pru-

minence is small and the dorsal protuberance is

nut detectable as a distinguishable area, The

dorsal prominence is on a Jevel with the

anterior rim of the acetabulum.

GLAUERTIA Loveridge

FIG. 3)

Speeies examinee: G. orientalis Parker,

The pubis and ischium are entirely car-

tilaginaus..

The. ilial shaft has an almost horizontal dor-

sal surface, and a sligntly curved ventral onc. so

creating a slight broadening ut the exuwemitics

of the shaft, There is neither « rim nor a crest

lo the shaft. The acetabulum has a s¢arcely

detectable peripheral rim which superiorly is

above the level of the ventral surface of the

ili) shaft. The ventral acetabular expansion ts

wlightly developed and the preacetabular zone

is slender, The dorsal acetabular expansion is

poorly developed. The dorsal prominence is

dificult to distinguish from the prominent and

evenly rounded dorsal protuberance which

uppears to be inclined laterally. Approximiately

one half of the length of the dorsal pro-

tuberance lics anterior to the anterior rim ol

the acetabulum,

HELEIOPORUS Gray

FIG. 3K

Speciey examined: H. albopunctatis Gray,

The pubis and ischium are ossified.

The tliat shaft is not curved but has o slightly

undulating superior face, raised inlo a

thickened fidge on the posterior half. The

PFLYIC GIRDLE, OF AUSTRALIAN FROGS 8

acetabulum is small and m surrounded by an

obliquely tapering rim. The ncefahulum is hish

and is bisectecdl by ihe veutral margin of the

ial shaft, The ventral acetabular expansion is

only stiehtty dilated but the dorsal acetabular

expansion rises sharply, The dorsal prominence

is Vast and the dorsal protuberunce simply an

elongate, semi-cylindrical ridge, At least two-

thirds of the dorsal protuberance lies anterior

to the anterior rim of the acetabulum,

KYARRANUS Moore

FIG, 32

Species examined: K. sphagnicolus Maore,

Rvarramniis sp.

The pubis and ischium are almost entirely

bony, only the central portion of the acetabu-

lar fossa remaining cartilaginous.

The ilial shaft ts strongly curved and almost

tircular in cross section, but for an indistinct

and narrow im on the posterior half. This tim

is created by an indentation of the medial sur-

face of the shaft, The ventral acetabular expan-

sion is poorly developed and the preacetabular

zone is very narrow, The dorsal acetabulur

expansion is quite prominent. The dorsal expan-

sjon is scarcely distinguishable from the Jarge

and oval, posteroventrally inclined dorsal pro-

minence. The anterior margin of the dorsal

prominence js considerably anterior to the an-

lerior rim of the acetabulum.

LECHRIODUS Boulenger

FIG. 3M

Species examined: L. melanopyga (Doria).

The pubis is cartilaginous and the ischium

ossified,

The ital shalt is slightly curved and bears

an enlarged, fanlike dorsal crest arising from

the posterior three-quarters of the shaft. The

acetabulum is small and has a broad peripheral

rim: the dorsal margin lies above the ventral

margin of the ilial shaft. The ventral acetabular

cxpansion is onty slightly developed. The dorsal

acetabular expansion is long and projects pas-

teriorly. The dorsal prominence is small but

detectable and the dorsal protuberance can be

distinguished, The anterior margin of the dorsal

protuberance is slightly posterior fo the anterior

rim of the acetabulum.

LIMNODYNASTES Fitzinger

FIG. 3N

Species examined: L. convexiusculus (Mac-

leav), FE, dumerili Peters, FL. ernatus (Gray),

L, peront (Dumesil & Bibron). L. sedmini Stein-

dachner, Z. spencer Parker, [ tseuamnfensis

Gunther, £, tevrueresinae Fry.

The pubis. is. cartilaginous and (te ischiam

is ossified.

The ilial shaft ts slightly curved and highly

vatisble in structure. There is an clongate

groove on the medial surface in L, dumerili, a

short groove in L. terraereginae, a short lateral

groove in L. rasneanternsis, and there is a dis-

tinct dorsal crest in L. ernates and ZL, spencert.

The acctabulum is small and high, brsected in

most species by the ventral margin of the ilial

shaft. The ventral acetabular expansion is small

and not particularly expanded. The dorsal

acetabular expanston rises into a moderate or

else high and acutely pointed spike. The dorsal

Prominence is conspicuous in all species except

those with crests on the shafls, Tr tends to fom

a conical shape in profile, but is somewhat

broader and rounded tn the largest species. The

dorsal protuberance is an elongate ndge or

expanded knob upon the tip of the dorsal pro-

minence. The dorsal protuberance is on a level

with or slightly anterior to the anterior rim of

the acetabulum.

MIXOPHYES Gunther

FIG, 30

Species examined: M. faselolatus Gunther.

The pubis and ischium are completely

ossified.

The ilial shatt is slightly curved and hears

an enlarged fan-dike dorsal crest arising dorso-

laterally from the Ienath of the shaft. The

acetabulum has a broad peripheral rim, the

dorsal margin of which lics above the ventral

margin of the ilial shaft. The vente) acelubu-

lar expansion is moderately developed bur with

a rather narow preacetabular zonc. The dorsal

acetabular expansion rises high to abil the

enlarged superior portion of the ischium. There

is no dorsal praminence, and the dorsal pro-

tuberance is entirely laterally directed and so

poorly developed that if is detectahle only when

the urea is viewecl from the dorsal or ventral

aspects. Whe anterior margin of this weak pro-

tuberance its located anterior to the anterjor rim

of the acelabulum.,

MYOBATRACHUS Schlegel

FIG. 4A

Species examined: M. gouldil (Grav),

The isthium is a large and almost circular

bony plate, and the pubis is reduced to a small,

triangular wedge of cartilage.

The iliwl shaft is distinctly curved, Jacks

crests und indentations und is slightly flatiened

10 M. J. TYLER

Fig. 4. Pelvis or isolated ilia of leptodactylid, microhylid and ranid frogs:

A. Myobatrachus gouldii, x 5; B. Netaden melanoscaphus, x 5; C. Neobatrachus centralis, x 5;

D, Philoria frosti, x 5; E, Pseudopkeyne bibroni, x 5; F. Ranidella parinsignifera, x 5; G. Rheo-

batrachus silus, x 5; H. Taudactylus diurnus, x 5; 1. Uperoleia sp., x 5; J. Cophixalus arnatus, x

5; K. Sphenophryne robusta, x 12.5; L. Rana papua, x 5,

laterally, producing an oval cross section. The

acetabulum is large and has a narrow

peripheral mm whose superior margin is con-

sidcrably above the level of the ventral surface

of the ilial shaft. The ventral acetabular expan-

sion is greatly reduced, consisting of just a

slender slip of bone bordering the acetabular

rim, The dorsal acetabular expansion is a more

prominent feature, rising above the ilial shaft.

There is no dorsal prominence, and the dorsal

protuberance is replaced by an oval, dorso-

laterally inclined plate, consisting of a weak

peripheral rim surrounding a very shallow

depression. The anterior margin of this struc-

ture is far posterior to the anterior margin of

the rim of the ucetabular fossa.

NEOBATRACHUS Peters

FIG. 4C

Species examined: N. centralis Patker.

The pubis is cartilaginous and the ischium is

ossified.

PELVIC GIRDLE OF AUSTRALIAN FROGS tl

‘The ilial shaft is almost perfectly straight.

The acetabulum is small and high, the ventral

marein of the jlitim being on a level with the

anterior une-third of the acetabulum. The ven-

iral acetabular expansion is only slishtly

dilated, the preacetabular zone being particu-

latly reduced. The dorsal acetabular expansion

is high. The dorsal prominence is very large

and the dorsal protuberance is a pointed nodule

upon it, Approximately one-half of the dorsal

protuberance lics anterior to the anterior rim

of the acetabulum,

NOTADEN Gunther

FIG. 4B

Species examined: N. melunoscaphus Hosmer.

The pubis and ischium are cartilaginous, the

laiter with a medisn calcified zone.

The ilial shatt is only very slightly curved,

lacks ridges and indentations and is circular in

eToss section, The acetabulum has a <listrnct,

flaltened peripheral rim which superiorly is on

a level with the ventral surface of the ifigl shaft.

The ventral acetabular expansion is slightly

developed with a narrow preacetahular zone.

The dorsal weetabular expansion is small and

modecratcly developed. The dorsal prominence

is broad and clearly demarcated from the ilial

shaft, The dorsal protuberance is small and

located on a level above the centre of the

acetabular fossa,

PHILORIA Spencer

FIG, 4D

Species examined: P. fresti Spencer.

The pubis is cartilaginous and the ischium

ossified,

The ilial shaft is sirongly curved and has a

medially directed dorsal rim, The acetabutim

is Large, its supenor margin on u, level with the

ventral margin of the ilial shaft. The ventral

acctabular expansion is not obviously dilated.

The dorsal acelubular expansion is moderately

well developed, The dorsal prominence is not

conspicuous and the dorsal protuberance

scarcely detectable as a separate entity. The

dorsal protuberance is almost. entirely antetinr

to the anterior rim of the acetabulum.

PSEUDOPHRYNE Filzinger

FIG. 4£

Species examined: P. bibront Gunther, P-

corfacea Keferstein

The pubis is cartilaginous and the ischitim is

ossified.

Phe ital shaft ls almost straighi and ts ¢ircu-

lar in Cross section, The acetabulum is large

and has a narraw peripheral tim, The superior

Margin of the acetabuluny lies on or slightly

above the level of the ventral margin of the

ijial shaft. The ventral acetabular exparision is

diated, The dorsa] acetabular expansion is only

very slightly raised. The dorsal prominence is

small (scarcely detectable in P. raridcea, but

quite distinct in P. bihfent) and the dorsil

prominence a very small knob on its tip. The

dorsal protuberance is on a level with the an-

terior rim of the acctabulum.

RANIDELLA Girard

FIG. 4F

Species examinedr R, parinsignifera (Main\,

®, sienifera Girard.

The pubis is cartilaginous and the ischium i3

bony.

The ilial shaft is curved, compressed Jaterally

and possesses neither a mm nor a crest. The

ucetabulum has a broad peripheral rim whicls

superiorly is on a level with the ventral margin

of the ilial shaft, The ventral acetabular expan.

sion is greatly enlarged, the subacetabular zone

protruding anteriorly. The dorsal acetabular

expansion is poorly developed. The dorsal

prominence jis broad and the dorsal pro-

tuberance is rounded, inelined posteroventrally

und moderately prominent, Approximately

one-half of the dorsal protuherunce is anterior

to the anterior rim of the acetabulum,

RHEOBATRACHUS Liem

FIG. 4G

Species exantined: R, situs Liem.

The pubis is cartilaginous and the ischium

ossified.

The iltal shaft is exceptionally slender, very

slightly curved and cylindrical in cross secon.

The acetahulum is very large with a con-

spicuous rim, its superior margin slightly above

the level of the ventral surface of the ilinl shaft,

The ventral acctabular expansion is slightly

dilated. The dorsal acetabular expansion is

slightly developed, projecting posteriorly. The

dorsal prominence is well developed and the

dorsal protuberance is conical and situated pos-

terior to the anterior rim of the acetabultm.

TAUDACTYLUS Straughan & Lee

FIG. 4H

Species examined: T, dikrnus Straughan & Lee

The pubis is cartifaginows and the ischium

is ussified.

The ilial shaft is gently curved, <tightly com-

pressed Interally and possesses neither a rim

2 MT. TYLER

nor g dorsal crest, The acetabulum is Jates with

a weil developed rim creating a deep acctibular

fossa, The ventral acetabular expansion is

slightly dilated. The dorsal acetabular expaiy-

sion is directed posteriorly and overlies the

superior margin of the ischium. The dorsal

prominence is only slightly distingnishable

from the large, raised, ovat, dorsolaterally

directed dorsal protuberance, Approximately

one-half of the dorsal protuberance lies anterior

toa the acetabular rim

UPEROLEI Gray

FIG. 4!

Species examined: U. marnworata Gray, LU pero-

leta sp,

The pubis and ischium are cartilaginous.

The ilial shaft is almost straight and is ctrew-

lar im cross section, lacking a rim and a crest.

The acetabulum is large and has a broad

peripheral tim, The superior margin of the

acetabulum lies above the ventral margin of the

ilial shafe. The ventral acetabular expansion is

of moderate width with the subacetahular zane

very slightly expanded. The dorsal aceiahilar

expansion is not pronounced. The dorsal pro-

minence ts large and the dorsal protuberance

conical and rising far above the level of the

shaft. The anterior margin of the dorsal pro-

tuherance is situated antenor to the anterior

rim of the aceiabulirm.

Family MICROIYLIDAE

COPHIXALUS Boetteer

FIG, 4/

Species examined: €. ornaus (Fry).

The ischium is small but ossified, and tre

pubis is cartilaginous,

The ilial shaft is compressed mediolaterully.

has neither a rim nor a crest, and is very

sheblly curved. The ventral acetabular expan-

sion is coticave and very narrow. The acetabu-

lar fossa is extremely large and very high. its

superior margin nearer to the darsal than the

ventral margin of che ilial shaft. The dorsal

protuberance is not distinguishable From the

dorsal prominence, and lies slightly anterior to

the anterior margin of the acetahular rim. The

dorsal acetabular cxpansion is very poorly

developed.

SPHENOPHRYNE Peters & Doria

FIG. 4K

Speciex examines; S, rohuyia (Yry),

The ischium is extremely small ind wnty

pany ossified. The pubis is cartilaginous.

The ilial shaft is compressed mediolaterally,

has neither a fim nor a crest, and is very

slightly curved, The yentral acetabular expan-

sion is slightly concave and very narrow. The

acetabular fossa is large and high, and is partly

bisected by the ventral border of the ilial shaft.

The dorsal protuberaice is not distinguishable

from the dorsal prominence, and lies entirely

posterior to the anterior margin of the acetabu-

lar rim. The dorsal acetabular expansion is only

slightly developed,

Family RANIDAE

RANA Linné

FIG. 4b

Species examined: R. papne Lesson

The pubis. is. cartilaginous and the ischium

is bony.

The ilial shaft curves gently downwards and

bears a tnassive fintike and tapering dorsal

crest, The acetabulum is large and bears a

broad petipheral rim. The superior margin of

the acetabular rim extends considerably above

the ventral margin of the ilial shaft. The ven-

tral acetabular expansion has a reduced pre-

acetabular zone and greatly dilated subacetabu-

lar zone. The dorsal acetabular expansion is

well developed, tapering to a point posteriarly.

There is no dorsal prominence and the dorsal

protuberance is an oval and almnst vertical

expansion of the base of the ilial shaft, This

profuberance lies entirely posterior to the an-

terior rim of the acetabulum,

Fossil genus

AUSTRALOBATRACHUS new genus

Type species: Australobatrachus ilius new

species,

This taxon was first reporied on by Tyler

(1974, p, 711, fiz. 1).

Extending from the acetabular region the

iial shaft bears o deep, curved groove an its

lateral surface. The acetabular fossa is excep-

lionally high in relation to the lial shaft. its

superior border reaching a position equivalent

to midway up the shaft. The acetabular cim is

poorly developed. The dorsal protuherance ts

gently rounded, and not distinguishable from

the dorsal prominence, The anterior limit of the

dorsal protuberance is on a Jevel with the an-

teriat margin of the acctabular rim, The ven-

tral acetabular expansion is neither protuberant

nor concave. the ilial/preacetabular zone form.

Ing almost a straight line extending gradually

posteriorly, The dorsal acetabular expansion

probably does not project superiarly,

PELVIC GIRDLE OF AUSTRALIAN FROGS 13

PIO ETE IT oe

| \| | il

yy)

WON Ni?

isi jf”

Fig. 5, Left ilium of Australahatrachus dins holotype. SAM, P18021.

Australobatrachus ilius new species

Aoelatype; Two tragments comprising the distal

6.7 mm of a single left tum. SAM, P1021

(Fiz, 5).

Type locatiry: Tedford Quarry, on the west side

of Lake Palankarrina, S.A. (Universily of

California Museum of Palagontology locality

V-5375.)

Horizon; Etadunna Formation.

Ase: Ngapakaldi Fauna, Tertiary; probably

mid-Miocene,

Deseription of holotype: As for gcnus.

Comparison with other species: Of the existing

families of frogs that do not have modern rep-

resentutives in Australia, the Leiopelmatidae.

Pipidae snd Pelobatidae have members in the

southern hemisphere and could all be regarded

as potential contributors ta the ancestral Aus-

Lralian frag fauna; hence they merit compari-

son with Australohatrachus.

The Letopelmatidae of New Zealand have

simple ila with a slender, cylindrical ilial shaft

and 4 poorly developed ventral acctabular

expansion (Stephenson 1960}. There is not the

slightest resernblance to Australohatrachys,

According to Trueb (1973), the pipids are

quite mnique in possessing a lateral crest (not

groove) to the ilial shaft. Any resemblance fo.

Aistralobatrachus has to be weighed against

the condition of the dorsal prominence (vast

and projecting high above the shaft in pipids;

reduced and not raised above the shaft in Amy-

trralobeltrachuy) and of the ventral acetabular

expansion (vestivial in pipids as oppased to

heing highly developed),

To judge from the descriptions and illustra-

tions of Zwetfel (1956) and Kluge (1966) the

pelobatid ilium has typically a bow-like dorsad

curvature, creating a totally different form from

that of Australobatrachus, In addition the

species illustrated by them have poorly-

developed ventral acctabular expansions and

more prominent acetabular fossae than has the

new genus,

Tn comparison with the ranids of Australia

and New Guinea, Australobatrachus is readily

distinguished by its total lack of the dorsal crest

which rises high above the ranid ilial shuft.

Similarly many microhylids (and also some lep-

todactylids) exhibit such a crest. but the micro-

hylids lacking a dorsal crest may he distin-

guished by the poor development of the ventral

acetabular expansion.

Rstablishing means of distinguishing Austra-

lian hylids Erom leptodactylids has proved diffi-

cult, In general it would apear that » well-

developed, dorsally-projecting dorsal pra-

minence is almost characteristic of Lhe lepto-

dactylids, whereas it tends 1!o be paorly

developed or else Jaterally disposed in hylids-

Most of the leptodactylids which are excep-

tional in having poorly-developed dorsal

prominences, are those in which the upper sec-

tion of the ilial shaft is modificd in some way-

Hence the prominence is scarcely differentiated

in Lechriodus and Mixophyes which have i

dorsal crest because the prominence is upon

this thin flange of bone. It could be argued

that the poor development of the dorsal pro-

minence in Australobatrachus conflicts with my

Interpretation of the groove as an intrinsic

modification of the ilial shaft. Hence an alter-

14 M, J. CYLER

native hypothesis is that the upper rim is the

supra-ilial structure, comparable to a dorsal

crest. This latter explanation is not favoured,

simply because the end section of the ilial shaft

iy S-shaped, and the nature of the acetabulum

(a poorly developed rim) is not usually

developed on a pelvic girdle in which the

acetabulum reaches the upper scgment of the

shaft.

Amongst the Australian hylids and Icptodac-.

tylids there is considerable variation in the

shape of the ventral acetabular expansion.

Cyclorana australiy and several species of

Litoria approach the condition displayed by

Anstrqlobatrachus, bat in each the dorsal pro-

minénce is more highly developed and the

lateral groove is lacking.

If Australobatrachus lacked a lateral groove

the nature of the dorsal prominence and the

ventral acetabular expansion would cause me

to favour referting the genus to the Hyliduc.

Hence to avoid over-interpreting the presence

and form of the Sateral groove, Australobat-

rachus is assigned tentatively to this family.

Extrapolation of musculature: The lateral sur-

face of the ilial shaft is the site of origin of the

musculus ifiacus externus. Any extreme

broadening of the lateral surface of the ilial

shaft increases the surface area available to

this muscle. Unquestionably these are madifica-

tions most Common in gquatic frogs or species

ifhabiting streamside situations. members of

such genera as Xenopus, Rana, Lechriodus and

Mixephyes, Certainly it is tempting to attribute

a similar functional association for Australo-

hatrachis, and hence assume that this animal

lived close to permanent water.

Acknowledgments

T am deeply indebted to Dr M, QO, Wood-

hurne for the opportunity to examine and des-

cribe the new fossil genus. Dr J, Ling and Mr

WN, Pledge read and provided constructive criti-

cisms of the manuscript, Mrs L. Kingston very

kindly typed the manuscript,

References

Cuantect, C. J. (1964).--Some Mio-Pliocene

hylids from the Valentine formation of Neb-

raska, Amer, Mid!. Nut. 72, 211-225,

Homan, J. A. (1965)—RFarly Miocene amirans

from Florida. Quart. J. Florida Acad. Sci. 28.

68-82.

Kutice. A. G. (1966)—A new pelobatine frog

from the Lower Miocene of South Dakota

with a discussion of the evolution of the

Scaphiopus-Spea complex. Contrih, Sei. Los

Angeley County Mus, (113), 1-26,

Lynen. J. D. (1963) —Additional evidence for the

recognition of Limnmaeodus (Amphibia:

Hylidae). Capria 1963, 566-568.

Lywen. J. D. (1971).—Evolvtionary relationships.

osteology, and zZeogeography of leptodactyloid

frogs. Misc, Publ. Univ, Kansay Muy. Nat.

Hist. (53), 1-238.

Srepumnson, E. M. (1960).—The skeletal churac-

ters of Lejopelia hatnilfani McCulloch, with

particular reference to the effecis of hetero-

chrony on the genus. Trans. Ro Sac, N.Z,

88(3), 473-488.

Trees, L. (1973)—Bones. frogs and evolution.

In Vial, J. (Ed.), “Evolutionary Biolagy of

the Anurans, Contemporary Research on

Major Problems”, pp. 65-132. (Univ. Mis-

souri Press.)

Trier, M. J. (1974).—First frog fossils from Aus-

tralia. Nature 248(5450),'711-712.

Zweirer, R. G. (1956)—Two new pelobatid frogs

from the Tertiary of North America and their

relationships to fossil and recent forms, 4m.

Mus. Novit. (1762), 1-45,

NEW AUSTRALIAN ALLODAPINE BEES (SUBGENUS EXONEURELLA

MICHENER) AND THEIR IMMATURES (HYMENOPTERA:

ANTHOPHORIDAE)

BY T. F. HOUSTON*

Summary

HOUSTON, T. F. (1976).-New Australian Allodapine bees (subgenus Exoneurella Michener) and

their immatures (Hymenoptera: Anthophoridae) . Trans. R. Soc. S. Aust. 100(1), 15-28,

28 February, 1976.

Three new species of Exoneura Smith (E. eremophila, E. setosa and E. tridentata) are described

and figured. They are assigned to the formerly monotypic subgenus Exoneurella Michener and both

adults and immatures of the new species are compared with those already described for the type-

species, E. lawsoni Rayment. A key for identification of adults is provided.

Females of E. tridentata vary greatly in size and exhibit allometric variation of the head and

metasoma.

NEW AUSTRALIAN ALLODAPINE BEES (SUBGENUS EXOQNEURELLA

MICHENER) AND THEIR IMMATURES (HYMENOPTERA:

ANTHOPHORIDAE)

by T. F. Houston*

Summary

Houston. T. F. (1976).—New Australian Allodapine bees (suhgenns Exonenrella Michener}

and their immatures (Hymenoptera: Anthophoridae). Trans. R. Soc. S. Aust, 100(1),

15-28, 28 February, 1976.

Three new species of Exoneure Smith (&. eremophilu, E. serosa and EF, widentata) are

described and figured. They are assigned to the formerly monetypic subgenus Exenenrella

Michener and both adults and immatures of the new spgcies are compared with those already

described for the fype-species, E. Jawsoni Rayment. A key for identification of adults is

provided,

Femules of E. midentafa vary greatly in size and exhibit allometcic variation af the head

and metasoma.

Introduction

The chief purpose of this paper is to provide

names for three species of bees. whose ethology

is ta be dealt with in a subsequent work, An

exhaustive examination of material from col-

lections has not been attempted und the des-

criptions to follow are based on specimens in

the collections of the author and the South

Australian Museum.

The three new species are assigned to the

subgenus Exoneurella Michener (of Exoneura

Smith) which formerly contained only the

type-species, E. Jawsoni Rayment. Since

Exoneurella was founded partly on the basis

of the larval characteristics of E. lawsoni, it

seemed of interest to describe and compare

immatures of the new species, Generally, the

characteristics of the new specics support re-

tention of Exoneurella as a discreet taxon.

The size-correlated variation of females of

E. tridentata sp. noy. is detailed below as’ it is

significant in terms of the bionomics of the

species. Such variation is unusval amongst

allodapine bees and in the family Antho-

phoridae as a whole.

The following abbreviations are used for

the names of institutions and collections te-

ferred to in the text below: ANIC ( Australian

National Insect Collection, C.S.LR.0O., Can-

berra), HE (author's private collection, to be

deposited in SAM), KU (Snow Entomological

Museum, University of Kansas. Lawrence,

Kansas, U,S:A,),; SAM (South Australian

Museum, Adefaide) and WADA (Western

Australian Department of Agriculture, Perth),

Except where stated otherwise, all specimens

listed in this paper were collected by the

author,

Genus EXONEULRA Smith

Exoneurd Smith, 1854, p. 232. See Michener.

1965, pp. 223-226 for detailed description,

subgenera and species.

Subgenus EXONEURELLA Michener

Exoneurella Michener, 1963, p. 257 (erected

asa genus); 1965, pp. 223-224 (relegated to

subgeneric status: diagnosis provided).

THE ADULTS

Key to the species of Exoneurella

1. Second cubital cell of fore wing with subequal

costal and medial borders (Fig. 1); 6th meta-

somal tergum of female with simple non-bific

apex and u pair of Jateral projections (Fig. 24,

25); compound cyes of male strongly swollen

(Figi8) -- gsm. a... 2 tridendia

1. Second cubital cell of fore wing with costal mar-

gin conspicuously shorter tham medial margin

(Figs 2. 3); 6th metasomal teraum of female

wiih bidentate apex and with or without lateral

* South Australian Museum, North Terrace, Adelaide, 5. Aust. 5000.

T. F. HOUSTON

FIGS 1-19

NEW AUSTRALIAN ALT ODAPINE HEES 17

prajections (Figs 4461, compound eyes of male

not swollen (Figs 17, |Bb - 1 2

2. Sixth metasomal tergum of fernale without

luteral prominénces, margins smonthly sinudle

(Fig, 4); hiod femora of male obttsely pro-

duced and curinule ventlully (Fig. &)) meta-

somal terga of both sexes black without

cream or ¢creamy-brown pigmentation and

withatl numerous cottspicuously thickened

setwe E. lawson

2. Sixth metasomal tergum of fernale with dis-

linct Joteral projections or prominences (Figs

5, 6); hind femora of male unmodified (Fig

7) or nal modified as in Fig. 8; mictasomal

tecea of both sexes with faint to distinct cream

bands; tergx 3 and 4 (females) or 4 and 5

(mates) with numerous conspicuously

thickened setae (Fig. 11) 0.0 0000... 9

. Metusomal terga yellow-brown to black with

wide cream bands (Pix. 14); 6th metasomal

tergum of female with subacute Jaleral projec-

tions (Fig. 6); hind femora of male cach with

an acute ventral projection (Fig. 9) 0...

E, eremophila

3, Metasomal terga Jargcly black with narrow

fyometimes faint or incomplete) cream subapi-

cal bands (Fig. 13); 6th metasomal tergum of

female with very obtuse Jaleral promiinences

(Fig, 5): bind femora of male unmodified (Fig.

os i oe , &. setosv

Exoncora (Exoneurella) eremophila n.sp,

Figs 2, 6, 9, 14, 15,17

Types

Holotype: &@ (SAM, 1 209613, New Kalamurini

Homestead, S. Aust. (27°44'8, 138°1S EB),

9-1 1.01.1972, on Wablenbersic.

Allotype: 2 in SAM, Poratypes; 42 d, 44 ° in

SAM:2 4,29 in ANIC, bf. 19 in KU.

Djacrosiy; This species differs [rom all other

Exzonenrella as follows. Lateral face marks of

male filling sptces between clypeus and com-

pound eyes (Fig. 17); metasomai terga of both

sexes yellow-brown to black with ‘extensive

cream maculations (Fig, 14): pronotam with a

pair of cream marks sublaterally; hind femora

of mule each with on acute ventral projection

(Fig. 9); hth metasomal lergom of female with

bidentate apex and a pair of subacuwre latcral

projections (Fig 4).

Deseriplion

Male. Body length 3.8-4.1 mm: head widih

1.]-1.2 mim,

Head capsule as broad as long; compound

eyes of usual relutive size (Pig. 17): face nur-

rowed to about 43% of head width; occlli

approximately equal in size to antennal sockets:

scapes failing to reach level of median occllus;

flagella 8596 as long as head width, the middle

segments about as long as broad: genae

viewed laterally almost half as wide as com-

pound eyes and evenly convex: fore legs not

especially slender, the Tore tibiae about 3.5x

us long as wide; bind femora laterally com-

pressed. cach with an acute ventral projection

(Fig. 9); costal margin of 2nd cubital cell of

fore wing about half as long as 1st transverse

cubital vein (Fig.2).

Integument glossy generally with sparse fitre

pitting or none; metasomal terga very finely

lineate,

Pubescence white, virtually absent dorsally

but dairly tong and dense ventrally and on legs

and mesepisterna;: 4th and 5th metasomal teres

fexcept laterally) with a sparse covering of

thickened bristle-tike setac.

‘the following areas white ar cream: lower

face, labrum and middle portions of mandibles

(Fig. J7). Seapes and pedicels venirally,

tubercles and dorsal margin of pronotum (ex-

cept medially), spots on tegulac, basal parts of

Wing veins, subapicul bands on tetusomul

terga (Fig, 14), apices of femora, bases of

tibiae, anterior edges of fore tibiae, and

basitarsi. The following areas yellow-brown:

scapes dorsally, flagella ventrally, most parts

of legs (except for cream areas), metssoma

Figs

i~ 3. Seoond cubilal cells of right fare wings (in dorsal views) of Evenewn (Exoneureila) triden-

tala, E. (B,) eremophila, and B. (F.) setesa respectively. C = costal margin, | = first

transverse cubital ven.

Figs 4~ 4, Sixth metasomal tergu (dorsal views) of females of £. (E.) la\vsoni, &. (E.) setosa and E.

(E.) eremophila respectively,

9. Trochanters, femora and apices of Ublue of teft bind legs (anterior views) of males of &

(E.) setesa, BE. (E,) lawsant-and #, (&.) erentaphila respectively

. Left mandible of female of E, (&) tridentata (ventral view).

. Thickened bristle-like sétae from fourth metasomal tergum of female of E. CE.) serosa

. Head capsule of a relatively large [emale of &. (&.) aridentaia (left lateral view),

and td. Metasomuae (dorsal views) of mules of 2. (2.) serosa und FE. 1.) erenemiila re

spectively showing cream bands (white) and translucent tergal margins (stippled)-

Fig. 15, Lower portion of head of female of E- (£.) eremophila (anterior view) showing T-shaped

chypeal mark,

Fig. tf

Head capsule of mete of 2, (E.) vidsmtatu (belt lateral view},

Figs 17-19, Head capsules (anterior views) of mules of RK. (fi) ercnnaptidl, BOR) setene and BL (E,)

(riglentofa fespectively-

\8 T. F, HOUSTON

ventrally and pastially or extensively dorsally,

Reinaiiioe areas Black or wack brown,

Vemale, Body length 4.0-5.5 mm: head width

1.i1-L2 mm.

Head form similar to thal of male (Fig. 17):

flagella about 67% as tong as head width;

mandibles tndentate but not constricted suh-

apically; metasoma fairly elongate, 6th meta-

sumal tergun with a bidentate apex and a pair

of small buat almose acure upturned lateral

projections (Fig, 6); hind femora unmodified,

Inlegument ane pubescence much as in male

hut thick bristle-like setae occur only on lergu

3 and 4,

Coloration as in mate except that while on

face ts limited to a full-length T-shaped mark

on elypeus (Fig. 15)

Pariation

The extent of yellow-brown coloration on

(he metasoma varies considerably amongst in-

dividuals collected together gnd some spect

menos have creamy-brown maculations on the

lateral margins of the scutum and scutellum.

The speertic piame, derived from Greek,

means loying solitude and alludes to the arid

hahitnt of the species.

Distribution

Central regions of Austealta including por-

tions of the Northern Territory, Queensland,

New South Wales and South Austratia.

Specimens examined: The holotype and the follaw-

ing. OLDS ¥ 3, 8 FP, A miles (4.8 km) W, of Wlo-

dara, U7,iv.1969, ex nests (HC), NWSI 1 2, 82

miles (232 kin) W. of Cobar, 9141971, on Prilo-

ts (SAM): | ct, 14 G70 miles (113 knit) E. af

Wileannia, 3111971, on Helichrysum, Goodenia

and Wahlenbergia (SAM). §. AUST.: | 3, Amata

(Musgrave Park) setuement, 14-%.1972, on Crlun-

drinta, H. BE. Evans & 1. P. Heuston (SAM): 1 9.

33 miles (33 km) W. of Amata, 17.x.1972, ex nest

in pithy twig, MW. &. Byany & T. FL Houston

(SAM); 7 &, Bettys Well (132°26'F, 2772’Sy-

Tiverund Park Sto, 1-S.xi.1970, on Aidiscies

fabrdgel (SAM 1; 43 ¢ (paratypes), 48 2 fallotype

and paratypes}, New Kalamurina HS, (27°44'S,

(38° 15'R), 9-01 71.1972, on Wahlenbergia (7 3,

5 F pinned), ex dead stems of Myriocephalins (38

¢, 43 Pin aleohol) (ANIC, KU, SAM): 2 &. Mor-

fat, 19.x11,1963, on fallenhereia (AC); 4 8, 49,

i} miles (16 km) S. of Mt Duvies airsirip,

71.%.1972, on Prlotis, ME, Evans & T. PF. Hons-

ior (SAM), 2 9 Mt Miceolle (32°31'S,

136°36E), Siam Stn, 2.iv.1971. ex nest (SAM);

1 9. 8. of Tomkinson Ranges (129°8°R, 26°10°S),

18.%.1972, on Sevevalu, H. E. Evans & 7. F, Hous-

ton (SANT): 2 f, 31 miles (40 km) W. of Wel-

bourn Hill HS, 13.%.1972. on blue Eremupliila,

HOE, EBvalis & TF, fhoustron (SAM): 2 2% 10

miles (16 km) SE of William Creck. 28,1,1277,

on Hakea, 1B Byuns & 7, F. Houston (SAM).

Froncura (Exoneurella) lawseni Raymeni,

1946, pp. 230-232, fig. 2 (male. not female or

Sarva)

FIGS 4, 8

Exenvaretla lawsont (Rayment)

1963, p. 257.

Franeura (Exoneurella}

Michener, 1968S, p. 224.

Holotype: 4 fin ANIC), Canberra, A,C.1..

Newlon R, Lawson. July (945,

1 hive not exammed the holotype ful Ms

Josephine Cardale (ANIC) made 4 critical

examinition of it on my behalf and confirmed

jhat Th agrees with the male churacleristics

given in the diagnosis below.

Michener (1963, p, 258) pointed out that

the fernales and larvae described hy Rayment

in the orginal description of E. Jawseni are af

unother species and are referable to. the sub-

genus Brevineura,

Diagnosis; E, lawsoni differs from all other

Exoneurella as. follows. Hind femora of mule

carinate and broadly produced ventrally (Fig.

8); 6th metasomal tergum of female lacking

lateral prominences, the margins gently sinuate

(Fig. 4). Differs from &. sétose and E.

cremophila in complete absence of cream pig-

mentation from metasoma and in wbsence (or

only feeble development) of thickened bristle-

like setae on dorsuni of metasoma,

For a detailed description of both sexes see

Michener (1963, p. 259). Note, however, that

the pale maculations of the face of the iiale

are white, not pale yellow,

Distribution: On and near Great Dividing

Range of south-eastern Queensland, New

South Wales and castem Victoria,

Specimens examined; OLD: 442,59, Bunya Moun-

tains, 6.x,1968, ex nesis THC). PIC. | fd, 2 3.

Tambo Valley, 221.1966, on Walilenheygla (AC),

Exoneura (Exoncarela) setosa n.s}),

FIGS 3.5, 7. IP. 13, 8

Michenes,

laweornt Rayment.

Types

Holotype: & (SAM, I 20962), West Beach,

Adetaide, S$. Aust, 25.iv,1975, ex deasl

Euphorbia stem, C. A. & T. F. Tfousten.

Allotrype: 2 in SAM, Parotypes; 7 ¢, 10 2 in

SAM; 42,4 2in ANIC; 23,2 9in KU.

Diagnosis: Very like E. lawsoni, differing a

follows. Hind femora of male unmodified,

lacking ventral flanges and projections (Fiz.

7); 6th metusomal tergum of female with a

pair of lateral convexities (Fig. 5); metasomal

terga of both sexes with narrow subapical

bands of creamish pigment (sometimes faint

or Incomplele, especially medially) and trans-

NEW AUSTRALIAN ALLODAPLNE BEES 19

lucent brown apical margins (Fig. 13): meta-

somal terya 4 and S (male) or 3 and 4 (fe-

male) With numerous conspictiously thickened,

hristle-like selac (Fig 11),

Dexeription

Male, Body length 3.84.7 mm: head width

1,1 1.2 mm,

Head capsule 11x as broad as longs com-

pound eyes of usual size (Fig, 18); face nar-

rowed ty about 42% of head width in lower

part; ocelli approximating size of antennal

sockets; scupes jusi failing to reach level of

median octellus) flagella about 724% as long as

head width; middle Mavellar segments slightiv

hroader than Jong; yenae viewed laterally 2/3

as wide as compound eyes and evenly conyea;

fore legs not unusually elongate. the fore tibiae

about 3.5x 4s long as wide; hind femora un-

modilicd (Pig. 7); custal margin of 2nd

cubital cell of fore wing about 1/4 ta 1/2 as

long as Ist transverse cubital vein (Fig, 3)-

Tntegument almost entirely glossy; clypeus

and scutellum finely pitted, dorsal area of pro-

podeunr dulled by extremely fine sculpturing;

metisumul terea finely lineate.

Pubescence white. sparse on head and body,

densest laterally and ventrally on thorax und

basal parts of legs: 4th and Sth melasomal

terga with numerous short but thick bristlyike

setae (Fig, 11) on dorso-apical areas.

The following areas white: almost all af cly-

peus and a spot of variable size on euch side

(Fig. 18), labrum, anterior stripe on Lore libia,

spots at bases of mid and hind tibiae, pronotal

tubercles and alar scleriles. The following arens

off-white to cream; ventral edges of scapes.

micl and hind basitarsi, narrow subapical bands

on metasomal terga (Fig. 13; sometimes. faint

or incomplete especially medially). Fore legs

(lurgely) and mid femora and tibiae anteriorly

yellow-brown. Hind margins of metusomal

ferga translucent pale brown; remaining reas

black or blackish brown,

Female, Body length 4,3-3.5 nim; head width

1.4-1.2 mm.

Head form similar to that of male: flagella

64% as long as head width; mandibles tri-

dentate but not constricted subapically:

melasoma elongate, the 6th lergum with bi-

dentate apex and w pyir af obtuse lateral

prominences (Fig, 3),

Inicgument sculptuled as in sale.

Pubescence much as in male but bristle-like

setae Occur on hind margin ef 3rd metasxomal

tergum and dorsal arga of 4th-

Coloration differs from that of male as fol-

lows: clypeus with a full-length white T-shaped

stripe; paraocular areas. without white spote;

labrum entirely brownish; legs lacking -yellow-

brown coloration.

The specific name, derived fron) Latin od

meaning *bristly’, alludes to the setation of the

rattasoma,

Distribunoa

Lowlands of southern South Australia (west

ta Spencer Gulf) and of south-ensternm Queens-

land.

Spectinens examined: The holowpe and the Follow-

ing, OLD: 1 3, 2 miles (3.2 km) S. of Nananeo,

7.x.1968, on Waltenberuia (HC), | 3 3 3 3

miles (4.8 km) N, of Peregian Beach {near Noosi

Ville), 9.xi1966, ex ness (HCO) S. AUST 2 dy

5.9, Glenely North (dunes), Adelaide, 24.%.1963,

€.x.1964 and 14-164,1965, on pigface and

Wahlenhergia flowers (IMC: 1 2, Mambray Creck

Rail Siding, 13.xi.1970_ ex pithy stem (SAM); 3 9,

Morgan, 18-19.xi7.1963, on Wahlenbergia (TIC):

13 df (paratypes), 17 2 (allotype and paratypes),

West Beach, Adelaide, same data as for holotype

(ANIC, KU, SAM): 4 3, 3 &, West Bench,

Adeloide, § and 24.11, 1965, 27.19.1965, on Wahlen-

bergia and ex bolfow stems (HO),

Kxoncura (Exoncurella) tridentata n. sp.

FIGS 1, 10, 12, 16, 19-30

Types

Holotype: J (SAM, F 20963), Lake Gilles

Notlonal Park (136°46°R, 33°2°S), S, Aust,

3) .Ai7.L973, ex short Linrel in twig of Meter

dendren.

Allotlype; 2 in SAM, Paratypes: 39 3, 29 9 in

SAM; 2 4.5 2in ANIC, 1 2,2 Yin KO.

Diagnosis: FE. tridentata differs fromm all other

Exonenrella as fallows. Coastal margin of 2nd

cubital eel] of fure wing equal to or slightly

longer than Ist transverse cubital vein (Fig,

1); scapes reaching to above level of median

ocellus; male with swollen compound eyes and

relatively narrow face (Pig. 19) and fore legs

conspicuously elonguled; female with angular

gennac (Fig. 12). mandibles constricted sub-

apically (Fig. 10) und 6th metasomal tergutn

with non bifid apex (Figs 24, 25).

Descripiien

Male. Body length +.5-4.9 mm: head width

1.6-1.9 mm.

Head capsule 7.3x us broud as long; comes

pound eyes strongly swollen so thal face ap-

pears sunken between them: face narrowed to

about 1/4 of head width ¢Pig. 19): ocelli

relatively large (about |.4x as wide as antennal

sockets}; genac viewed laterally (Fig. 16)

much narrower than compound eyes and nal

angular; scapes slender, reaching to just above

20 T. F, HOUSTON

Tmm

tnt

Figs 20-29. Fxoneura (Expneurella) tridentata female. Figs 20, 21.—Heuad capsules (anterior views) of

smallest and largest known specimens respectively, drawn to same length, Figs 22, 23—

Smallest and largest known specimens respectively (dorsal views) (antennae and legs omit-

ted, fore wings represented by broken lines). Figs 24, 25,—Sixth metasomal terga (dorsal

views) of smallest and largest known specimens, respectively. Figs 26, 27—Teft halves of

fourth metasomal sterna (dorsal views) of smallest and largest known specimens, respec-

tively, drawn to same length, Figs 28, 29.—First metusomal terga (dorsal views) of smallest

and largest known females, respectively, drawn to same size to illustrate differences in sur-

face pitting.

level of median ocellus; Hagella relatively short,

L/2 as long as head width, all segments but

apical one broader than Jong; fore lees very

slender, the tibiae 5x longer than wide; hind

femora unmodified; costal margin of 2nd

cubital cell of fore wing equal fo or longer

than Ist transverse cubital vein (Fig. 1);

tmetasoma relatively short, broad and de-

pressed,

Integument, of face smooth but dull with

close small pitting on clypeus; scutum and

scutellum glossy with very sparse fine pitting;

mesepisterna dulled by shullow coarse pitting;

anterior half of Ist metasomal icrgum glossy

and impunctate. the posterior half and most of

tergum. 2 pitted and finely roughened, dull ¢x-

cept laterally; tergum 3 shiny but coarsely

Pilled; terga 4 to 7 duller with fine reticulate

sculpture.

Pubescence white, fairly long and sparse

generally, densest on clypeus, posterior of

head, sides and venter of thorax, sides of pro-

podeum, basal areas of legs and ist metasomal

tergum,

Colour black generally except for the follow-

ing: clypeal mark (Fig. 19), ventral edges of

scapes, patches on tegulae and wing bases and

spot at base of each tibia white to cream;

apical portions of femora, all tibiae and tarsi

orange-brown; wing veins and ventral surfaces

of flagella brown.

NEW AUSTRALIAN ALLODAPINE REES 21

Memale. Size eatremely variable: bedy length

$,5-10.0 mm; head width 13-2.) mm.

Head (viewed aliteriotly) grading from

fairly round in small femules (Fig, 20) to

rather quadrate in large females (Fig. 21);

uccordingly whe inner orbits yary from slightly

converging to slighily diverging below. genae

(viewed laterully) almost as wide as compound

eyes and very angular, especially m large

specimens (Fig. 12); scapes slender and reach-

ing median ocellust Aagella 659 as long as

hend width; labrum) with a stout carinate

median tubercle; mandibles tridentate, strongly

constricted subuapically (Fig, 10); legs not as

slender as those of male; metasoma elongate

and rather parallel-sided, more so in large f{e-

males (Figs 22. 23); 6th tergum upturned,

slightly to strongly concave dorsally. rather

Iriangular with small Jateral projections in

small females (Fig. 24) grading to quadrate

im Jarge females (Fig. 25),

Totegument largely glossy with few scattered

small pits; dorsal and lateral arcas of pro-

podem dull with fine roughening; Ist mela-

somal tergum of small specimens with re-

latively few pits conogntrated along posterior

margin (Fig. 28), of larger specimens with

numerous coatser pits some of which exceed

the ocelli in size und many of which coalesce

to form at irregular emargination posteriorly

(Fig. 29), more apical terga with sparse

niedium pitting and fine reticulate sculpturing,

sironges! on terga 4 jo 6. Pubescence generally

sparse, white and inconspicuous, longest on

sides of metasoma and hind tibiae; dorsal arcas

of metasoinal terga lacking thickened or con-

Spicuous setac,

Head and body black) Glypeus with a full

length T-shaped white mark: labrum, man-

dibles and leys largely or wholly orange-hrown;

medium and Jarge females usually have diffuse

oranze-brown patches on mesepisterna, meta-

sternum and anterior metasomal sterna.

The specific name refers ta the 3-pointed

margin of the 6th metasoinal tergum of the

female.

Disrriburian

Semrariu rogians of Suuth Australia ond

southern Western Australia (the mallee Enca-

Iyptus belt und bordering areas),

Specimens examined: The holotype and Lhe faltew

ing. S AUST: 1 # and 14 2 (Coll paratypes).

Corunna Hills, N. of Tron Knob, 19.4v.1971, ex

nest (SAM); 4 9, 8. of tron Baton, Byre Penin-

sula, 30x,1971, on Eremoplile (HC); 40

(paratypes}, 26 2 {including allotype and 22 para-

types}, Lake Gilles National Park, 30-31..0.1973,

TL=0fRiV.9974, b4-17.Vi,1974, 29.viii-1.ix, 1974 and

27-x.1974, ox nests in hollow Hererudendren lwigs,

©. A. aT. &. Huuston (ANIC, HC, pi und SAM

—some in alcohol); 1 (paratype). 3 9, norlhera

Middleback Rungey (137°9'E, 33°3'S), 7-8.x. 1978,

ex hollow Meteradendrim twigs, CG, Ao & TT. B-.

Houston (AC, SAM), 7 9. 8 miles (13 tum) E. of

Pecener, 8.11970, on, Melaleuca pabescens (HC);

L ?, 29 km NNW, of Pt Augusty, 29.1%, dae on

Myaperum (SAM), 3 9, 2 miles (3.2 km) N, of

Port Germejn, 7.7,1970, on Lorawithns niraculosns

and Mefaleucu pubescens (UIC); 1 3. 30 miles (48

km) NNW. of Renmurk, 22.1, 1972, on mallee

Eucalypius (SAM), W,. AEST. LS, 27 miles (43

km) W. of Coolgardie, 38.1,1970, on Evendy pine

(HC): 1 9, 25 miles (40 km) EB. of Kalbarti,

BAITS. KT. Richards Hef aad b | 2 & miles

(13 km) S. of Wanvo, 7-v-1974, TY, Richards

(WADA),

Size-correlated variation in B. tridentata

As ioted in the above description. females

of &. tridemare vary miurkedly in size and form

and slightly in coloration. Same of this varia-

tion is correluted with size and since it is sig-

nificant in terms of the bionomics of ihe bees

it is detailed below.

Individual size has been judged according ta

head ¢apsule width. Measurements of fore

wing. lengths wete made but proved un-

necessary since the ratio of fore wing length to

head width remained approximately constant

OVer a range of mensurements.

With increasing size, the following changes

occur.

(1) The head capsule becomes increasingly

more quadrate, the inner orbits changing

from slightly converging to slightly di-

verging below and the clypeus becoming

wider relative to its length ¢cf, Figs 20,

21).

The metasoma becomes felitively larger

(especially in length}, In the smallest fe-

males the apex of the metasuma does not

extend beyond the tips of the reflexed fore

wings whereas in the largest females it

protrudes beyond them by about 1/3 of

its length (cf. Figs 22, 23). This increase

in relative size is mot due to extension of

the telescopic segments but reflects an in-

crease in size of all the component parts

including the sting.

The latero-apical projections of the 6th

mictasomal terguy become relatively

larger, more obtuse and further apart (cf-

Fins 24, 25). This variatian is quantified

in Figure 39.

The Ist metasomal tergum hecrmes in-

creasingly more coarsely and deeply pitted

and in the Jarger females the pits slong

(4)

Apuen) width of loth, mérasomot teeauen 6 Meme wldih (7)

a té 1.8 zo 22

Head widrh trem}

Fip, 30. Allometry in females of Exenenra (Exo-

Neuretle) tridentata. Seatter diagram

showing how relative width of apex of

sixth metisomwl tergum increases with

size of individual. The umber of indivi-

duals répresented hy spots increases with

their size in the order J, 2, 3, 4, 5, 6-10,

11-45, 16-20 and 21-25,

the posterior margin coalesce to form an

ittegular emargination (cf. Figs 28, 29).

The integument of the metasoma becomes

relatively thicker and more brittle and the

apodemes become relatively larger (cf.

Figs 26, 27). The sturdier anterolateral

apodemes are associated with relatively

lurger extensor muscles.

Orange-brown patches with diffuse bor-

ders appear on the thorax and metasoma

of medium and lurge females, They occur

on the mesepisterna, mesosterna and

anterior metasomal sterna. The larger the

individual, the more extensive are the

maculations.

A small median spine develops from the

gradulus of the 4th metasomal tergum in

medium and large females.

ti}

CO)

(7)

T. F. HOUSTON

This variation appears to be unique amongs}

allodapine bevs und in the family Antho-

phoridac as a whole, Michener (1965a) dis-

cussed size variation amongst females of the

social] Australian hee, Exenevra (E.) yariabilis

Rayment, In this species, egg layers average

larger than workers but na structural differ-

ences or allometry has been reported. The

degree of size variation in this species is also

less than in &. tridentata, the largest known

females having head widths only 1.28x as great

as the smallest females (cf. 1.54x in E,

iridentata)

THE IMMATURES

The immatures studied wore preserved by

dropping them live into either Kahle’s solution

or 75% ethyl alcohol and were stored in the

latter,

Syed (1963) described 4 larval instars of

EY lawsont from preserved material and

Michencr (1964) described and figured live

specimens of probable 2nd, 3rd and 4th instars

as well as the egg, prepupa and pupa. How-

ever, the identity of the material studied and

described by these wuthors is subject to a little

uncertainty hecuuse a few of the nests from

which it was devived have proven to belony to

EE. setosa (C,. D. Michener—personal com-

munication), Professor Michener confirms thit

the bulk of the adult material on which he

based his 1964 studies is clearly E. Jawsoni and

in all probability the immatures described by

him would be of the same species,

Four morphologically distinct larval instars

can be recognised in each of the known species

of Exoneurella without recourse to histograms

of head width frequencies. Indeed, with

Exoneura tridentata there is such marked size

variation, within each instar and such wide

everlap in size between them that the histo-

gram of head width frequencies was of no

help at all in determining the number of in-

stars.

The terminology employed in the following

descriptions of liurvag follows that of

Michener (1953).

Figs 31-42. Immatures of Exoueura (Exoneurella) eremophila. Fig. 31—Ege. Fig. 32.—First instur

partly enclosed in chorion (left laterul view). Fig. 33-—Second instar with chorion still

attached (left lateral view), Fig. 34,—Ventral view of head and fore body of secand in-

star. Figs 35, 36.—Third and fourth instars, respectively (left lateral views). Fig. 37.—

spiracle of fourth instar, Fig. 38—MNead of fourth instar (anterior view). Figs 39, 40.—1 eft

mandible of fourth instar, anterior and ventral views, respectively. Fig, 41.—Mouthpasts

of fourth inslar (posterior view). Fig. 42.—Female pupa (left lateral view).

NEW AUSTRALIAN ALLODAPINE BEES

FIGS 31-42

24 T. F. HOUSTON

E. (E.) eremophila

Beg, (Fig. 31). About 4.1 mm: long and 0.4

mm in maximum width; white, sausave-shoped

and with a coarse Teticulate sculpturing each

end, the middle portion being tuberculate,

In some eggs from Queensland nests the

sculpturing was weak or absent.

first instar—(Fig, 32), Remains almost

wholly withitt chorion, head of very simple

form, lacking lateral lobes, antennae and setac;

mouthparts hardly developed, lobe-like and

probably non-functional: body sac-like without

obvious segmental lines, tubercles or sctac.

Second instar—tFigs 33, 34), Chorion re-

mains attached to abdomen, head selatively

broader than in |st instar but with no obvious

lateral lobes; antennae well-developed, capitate

and laterally directed; mouthparts well-deve-

loped and functional: bead capsule wilh

numerous moderately long setae; body without

setae und distinct segmental lines but with 4

tubercles: each side anteriorly and a middorsal

tubercle on prothorax.

Third instar— (Fig. 35), Entirely free of

chorion; head relatively very broad with well

developed ventrolateral lobes; antennae yery

slender and acute apically; body gently curved

with distinct intersegmental lines, no antero-

lateral tubercles or ventrolateral swellings but

prothorax with a distinct middorsal rubercle;

anal slit moderately deeply incised; sctac

numerous on head and body, longest on

ventrolateral lobes of heady patches of small

setae occur dorsally and laterally on the pro-

thorax, laferally an the following 4 segments

und transverse rows of shart stiff setae occur

dotsolatcrally on the 2nd to 12th badly seeg-

ments,

Fourth instar —(Fivs 36-41). Head relatively

very broad with conspicuous ventrolateral

swellings and slender acute antennae (Fig. 38);

labrum bilobed apically with a few sensoria,

not delimited from distinctly sunken clypeal

region: mandibles slender apically with only a

few minute spines. subapically and a single

sensorium ventrally (Figs 39, 40): manxillae

shorter than labium, their palpi consisting of

indistinct tubercles bearing a few sensoria

(Fiz, 41); labium bearing tubercle-like palpi

latero-apically; body strongly bent at Sih ab-

dominal segment (Fig. 36); prothoras with an

obtuse mid-dorgal tubercle, intersegmental lines

wenk: ventrolpteral body swellings ubsent;

terminal segment of abdomen strongly laterally

compressed With anal slit deeply incised; setn-

tion much as in 3rd instar but all holy seg-

ments have ventrolateral patches of sctac;

atrial aod) piimaty tracheal openings of

spiracles circular; atria without spines but with

a Jew branching and anasloumosing seulpiural

lines; subatria relatively Jong (Fig. 37}.

Prepupa,—Similar to 4th instar except that the

body is straight and swollen anteriorly.

Pupa—{Fig. 42). Conforms essentially to

features of the adult bur the following spectul

pupal structures were noticed: all coxae with

ventro-apical spines (very short and incgon-

spicuous un mid and hind coxae of femules);

vertex (across full width), interantennal area,

upper, middle and lower clypeus with ea-

tremely long setae; 2nd to Sth metasomal seg-

ments also with 2 or 3 lonw setae each side; in

same speciinens the more apical metasomal

terga bear u few tiny setae dorsally,

Material examined.--155 eggs, LOY larvae and pre-

pupac and 46 pupae, New Kalamurina HS, $8.

Aust., SQ ILUL1972, ea dead stems of Myyia-

cephalus; 7 eggs, 48 larvae and prepupac and 2

pupue, 4.8 hm W of Windorah, Qld, 17.14.1969, ex

dead stems of Crotalaria,

FE, (F,) setosa

Eey.—( Fig. 43). About 1.0 mm long and

0.4 mom in maximum width: white, sausage-

shaped and with 4 reticulate sculpourul pattem

(finer than that of £. ereinophila eggs, }

First instar —Nol observed,

Second invstar-—( Pig. 44). Similav te that of EB.

eremophila except that antennae pre not eapi-

tale; Ist and 2nd body segments with a few

small setac dorsally.

Third iusjar—{ Fig, 45). Similiar 10 thot of BL

eremophila hor with more distinct, interseg-

mental Jines and moderately developed ventro-

luteral body swellings. Of 4 specimens ex-

amined, 2 lacked dorsal abdominal setae and

2 slightly Tarver ones had setae on all but the

terminal segment,

Fourth iastar—(Pig. 46). Generally similar

to that of E. eremophila except as follows:

intersegmenial lines more distinct; ventro-

lateral swellings moderately developed; Sth

abdominal segment slightly more proluhetant

dorsally; labial palpi situated more posteriorly

on labiumy and further from apex,

Prepupa—Like 4th instar but body straight

and swollen anieriorty.

Pujt—Generally similar ta thar of &. eres

phile (allowing for differences correlated with

NEW AUSTRALIAN ALLODAPINE HEES a5

1 mm

Wigs 43-46. Immatures of Exonéarg (Bxo-

Nenrella) sétasq. Fig, 43.— Egg, Fig.

44.—Second instar in left lateral

(upper) and dorsal (lower) views.

Figs 45, 46.—Third and fourth in-

stars, respectively (left Jateral views),

adult form); 3rd tq 5th metasomal terga with

a few short setae dorso-apically.

Material examined—21 eges, 32 larvae and pre-

pupae und 12 pupae, West Beach, Adelaide, S.

Aust,. Feb.—Oct. 1965, ex dead stems of Exphor-

bia; 14 eggs, 28 larvae and prepupae and 3 pupue,

4.8 km N. of Peregian Beach, Qld, 9.xii.1966.

E. (E.) tridentata

Ege.—(Fig. 47), Size variable ranging from

1.3-2.0 mm in length and 0.43~0.70 mm in

maximum width; white, sausage-shaped ta

elongate ovoid; chorion entirely smooth to

finely granular (except at the ends),

First instar —(Fig. 48). Remains largely with-

in chorion; head capsule smooth and appoxi-

mately circular in anterior vicw; antennae,

mouthparts and setae absent; body sac-like

without intersegmental lines, tubercles and

setae.

Second insiar—(Fig, 49), Retains chorion on

apex of abdomen: head broad with distinct

lateral lobes which are usually reflexed against

sides of prothorax; antennae absent; mouth-

parts developed and functional: body sausage-

shaped, curved, lacking tubercles and with

weak intersegmental lines; head with numerous

setae, longest on vertex where they are thick

throughout their length; body lacking setae.

Third instar—(Fig. 50). Head rélatively ex-

tremely broad with laterally extended lobes;

untennac present, relatively short (compared

with those of other Exoneurella) and medially

directed; body curved. without tubercles

{except dorsa-apically on terminal segment)

and with weak intersegmental lines; head with

almost a complete covering of short blunt

setae; body without setae.

Fourth instar—(Figs 51-56). Head with ex-

ceptionally large quadrate lateral extensions

(Fig. 53); antennae short, slender, acute and

medially directed; labrum broad und bilobed

with several sensoria ventrally (Fig. 56), not

delimited [rom clypeo-frontal area: mandibles

(Pigs 54-56) slender, tapering and compressed

apically, cach with a pair of sensoria ventrally

but lacking spines; labium rounded and lobe-

like, bearing 2 patches of sensoria which may

represent obsolescent palpi (Fig. 56): a trans-

verse, laterally projecting lobe behind the

labium probably represents degenerate maxillae

(Figs 53, 56): bedy strongly bent at Sth ab-

dominal segment which, like the 6th, bears a

prominent dorso-median tubercle (Fig, 51);

7th ahdominal segment with a small dorsal

tubercle; terminal abdominal segment broad

26 T. F, HOUSTON

Figs 47-56. Immatures of Exoneura (Exoneurella) tridentata. Fig. 47.—Eggs showing extremes of form

and sculpturing. Figs 48, 49.—First and second instars with chorion attached (left lateral

views). Figs 50, 51.—Third and fourth instars, respectively (left jateral views), with en-

largements of capitate seta and spinose apical tubercle. Fig. 52.—Spiracle of fourth instar.

Fig. 53—Head capsule of fourth instar (anterior view). Figs 54, 55.—Left mandible of

fourth instar in anterior and ventral views, respectively. Fig. 56—Mouth parts of fourth in-

star (ventral view. L, labrum; LI, labium; M, mandible; MX, maxilla?). Figs 48-51 are

drawn to same scale.

NEW AUSTRALIAN ALLODAPINE BERS 27

with a moderaicly Jarge spinose tubercle just

above the anus (Fiz. 51); head with mmerous

short obtuse seige: body with short setue dis-

posed in tramsverse bands on prothoracic to

4th abdominal seaments and small dorsal

patches on the 5th to 9th! relatively large thick

capitate selac occur ventrally in clusters on the

metathoracic to 2nd abdominal segments and

singly or clustered Iluterully on the meso-

thoracic 10 4th iabiulominal sezments; spiracles

(Fig, 52) nol protruding above body wall;

autal ark primary tracheal openings circular;

aria subspheroidal without spines or other

sculpture, subglria slender with about 18 an-

nulations.

Prepupo—similar to 4th instar but body

straight, swollen anteriorly and with reduced

dorsal tubercles,

Pupa—Similar to that of &. eremoplila dif-

fering as follows. Head with fewer setae, a puir

being silualed tow on the clypeus and several

across the vertex. the more lateral ones being

much longer than more medial ones; metasoma

with short setue dorsally on segments 2-5 (fe-

mates) and 2-6 (males) fin addition to the

long lateral setae).

Material examined. 2004- eges, 125 latvae and

Prepupac snd 73 pupae, Lake Gilles National

Park, S. Aust, 30,x01.1973-27,..1974, ex desd

slems of Helerodendion olelfolinn.

Discussion

Regarding adult features, the 3 new species

agree almost lotally with the diagnosis of

Frorenrella given by Michener (19654, p.

223). The only points of disagreement relate

io E. tridentata) in this species the eyes of

males are conspicuously swollen, the apex of

the 6th metasomal tergum of females is simple.

net bifid, and the costal margin of the 2nd

cubital cell of the fore wing of buth sexes is

at least as Jong as the Jat tramyverse cubital

vein, not much shorier, In these respects E.

teidentata is rather more like bees of dhe sub-

genus Exoneura than are other Exonenrella. It

is also unlike its closest relatives in the con.

stricted mandibles, angular genac, pronounced

size variation and allometry of females.

Eggs of Ezoreurelia are wnusual amongst

those of allodapines {Michener 1973, p. 281)

in having sculptured chortons, The sculpturing

forms a delicate reticulum in E, lawsent and

E. velosa, fine sranules in &. iridermata (absent

in some specimens) and a cumbination of

coarse reliculum and distinct tubercles in A-

ereniophita.

Similaritics in larval [orm between the 4

species of Exonenrelie correspond to similar.

ties In adult form. hus, larvac of BE. serosa

are most like larvac of &. fawsoni as described

and figured by Michener (1964, pp. 422-424,

gs 13-20) and larvae of E. eremaphila differ

from these 2 species in only a few minor

features. On the other hand, larvar of &,

tridenjata are highly distinctive: the head cap-

sule of 2nd to 4th instars is extraordinarily

produced and quadrate laterally, the antennae

are comparatively liny, the maxillae and

labium are strongly modified, 2nd instars lack

Jateral body tubercles but 41h instars have large

dorsal tubercles on the 5th and 6th abdonnnal

segments, spiny apical tubercles and peculiar

thickened setae on the thoracic and anterior

abdominal segments.

The fentures which will distinguish

Exoneurelia larvae from those of other groups

acc the following. Head capsule fof more

mattire instars) relatively broad with distinet

hairy ventrolateral of lateral expansions; an-

tennae (except in £. rrideniata) of 2nd to 4th

instars relatively long, slender apically, thick-

ened basally and directed anterolaterally:; na

separuhon of clypeus and Jabrum; mandibles

strongly tapered with slender simple apices;

body of 4th instar conspicuously bent at Sth

abdomina) segment which protnides dorsally:

3rd and 4th instars lacking lateral or ventro-

lateral extensions of hody segments such ag

occur in other Exoneura (Syed 1963),

The pupse of Fxoneurella differ from

species to species in conformity with adult dif-

ferences bul otherwise are fairly uniform. Of

the various specialized pupal structures occur-

nng in Apoidea (Michener 1954) the only ones

occuring in Exoneurella ute lang thick setae

on the head and mectasoma, fine shost setae on

the metasomal terga, and coxal spines

Michener (1964, p. 424) remarks on the ab-

sence of coxal and trochanteral spines in E.

fawsen? but I have seen no material which

could confirm this. Specific differences were

noted in the number and arrangement of setae,

Exoneurella. originally established as a genus,

was felegated to subgeneric§ status in

Michener’s (19655) classification of Australian

bees but has continued to receive generic

status (Michencr 1971, 1973). ‘The taxon with

ils new additions remains distinctive and well

defined, T consider it & pulely arbitrary malter

whether one recognizes it at generic or sub-

generic Icevel and have preferred ta Inllow

28 T,.. F. HOUSTON

Michener’s (19656) arrangement since it ex-

presses the obvious affinity between Exo-

neurella, Exoneura s. str. and Brevineura.

Acknowledgments

I wish to thank my wife, Carol Houston, for

assisting with collection of some of the

material described below, Ms J. C. Cardale

(ANIC) for providing some critical details of

the type specimen of Exoneura lawsoni, and

Professor C. D. Michener (KU) for advice

concerning material used in his studies of

Exoneurella.

Some of the material listed herein was col-

lected during field work funded by the Royal

Society of South Australia.

References

MICHENER, C, D. (1953),—Comparative morpho-

logical and systematic studies of bee larvae

with a key ta the families of hymenopterous

Jarvac. Univ. Kans, Sci. Bull, 35(8), 987-

1102.

Micnener, C. D. (1954).—Observations on the

pupae of bees (Hymenoptera: Apoidea). Pan-

Pacif. Ent, 30(1), 63-70.

MIcHENER, C. D. (1963),.—New Ceratinini from

Australia (Hymenoptera, Apoidea). Uniy.

Kans. Sci, Bull. 49(7), 257-261.

Micuener, C, D, (1964).—The bionomics of Exo-

neurella, a solitary relative of Exeneura

(Hymenoptera: Apoidea: Ceratinini). Pacif.

Insects 6(3), 411-426.

MICHENER, C. D. ([965a).—The life cycle and

social organization of bees of the genus

Exoneura and its parasite, Jnguilina (Hymen-

optera; Xylocopidae). Univ. Kans. Sci. Bull.

46(9), 335-376.

MICHENER, C. D. (1965b).—A classification of the

bees of ihe Australian and South Pacific

regions, Bull, Am. Mus. nat. Fist. 130, 1-362.

MicHener, C. D. (1971).—Biologies of African

allodapine bees (Hymenoptera, Xylocopinae).

Bull, Am. Mus. nat. Hist. 145(3), 219-302.

Micuener, C. D. (1973).—Size and form of eggs

of allodapine bees. J. ent. Soe. sth Afr. 36(2),

281-285.

RAYMENT, T. (1946).—New bees and wasps—

Part Il. Victorian Nat. 62, 230-236.

SmitH, F, (1854) —“Catalogue of hymenopterous

insects in the collection of the British

Museum. II.” (British Museum: London.)

Syrp, I. H. (1963).—Comparative studies of lar-

yae of Australian ceratinine bees (Hymenop-

tera, Apoidea). Univ. Kans. Sci, Bull, 44(8),

263-280.

FAULTING CONTEMPORANEOUS WITH UMBERATANA GROUP

SEDIMENTATION (LATE PRECAMBRIAN), SOUTHERN FLINDERS

RANGES, SOUTH AUSTRALIA

BY P. §. PLUMMER* AND V. A, GOSTIN*

Summary

PLUMMER, P. S., & GOSTIN, V. A. (1976).-Faulting contemporaneous with Umberatana Group

sedimentation (Late Precambrian), southern Flinders Ranges, South Australia. Trans. R. Soc, S.

Aust. 100(1), 29-37, 28 February, 1976.

Interglacial sedimentation within the Late Precambrian Umberatana Group (Adelaide System) was

greatly influenced locally by contemporaneous faulting along the Spring Creek Mine Fault.

Stratigraphic study both sides of the fault and of the fault zone has revealed a variable sequence of

elastics and carbonates. Lithological correlation across the fault zone suggests two periods of

faulting contemporaneous with deposition, followed by a phase of downwarping of the area north of

the fault along a hinge line coinciding with the earlier fault zone. This activity resulted in a

thickness increase of approximately 750 m in the sequence north of the fault when compared with

that of the more stable sequence to the south. Three varieties of stromatolites occur within the area

studied, displaying distributions which were influenced by varying water depths.

Palacoenvironmental interpretations based on vertical and lateral lithological associations,

palaeocurrent analysis and studies of the contained stromatolites reveal that sedimentation south of

the fault occurred within shallow marine, intertidal and supratidal environments resulting from the

prevailing regional regressive-transgressive-regressive marine cycle. Modification of this cycle

north of the fault zone was influenced by contemporaneous faulting.

FAULTING CONTEMPORANEOUS WITH UMBERATANA GROUP

SEDIMENTATION (LATE PRECAMBRIAN), SOUTHERN

FLINDERS RANGES, SOUTH AUSTRALIA

by P. 8. PLuuMMeR® and V. A. Guostin*

Summary

PeumMer, PF, S. & Gostin, V, A. (1976).—Faulting contemporaneous with Umberatana

Group sedimentation (Late Precambrian), southern Flinders Ranges, South Australia,

Trans. R. Soe. §. Aust. 100¢1), 29-37, 28 February, 1976,

Intergiacial sedimentation within the Late Precambrian Umberatuna Group (Adelaide

System) was greatly influenced locally by comemporaneous faulting aluoe the Spring Creek

Mine Fault, Stratigraphic study both sides of the fault and of the fault zone has revealed a

variable sequence of clastics and carbonates, Lithological correlation across the fault zone

suggests two periods of faulting contemporaneous with deposition, followed by a phase: of

downwarping of the area north of the fault along a hinge line coinciding with the earlier fault

xone, This activity resulled in a thickness. increase of approximately 750 mw in the sequence

north of the fault when cornpared with that of the more stable sequence to the south. Three

varieties af stromatolites occur within the area studied, displaying distributions which were in-

Nuenced by varying wuler depths.

Palaeoenvironmental interpretations based on vertical and lateral lithological associations,

palaegcurrent analysis and studies of the contained stromatolites revenl that sedimentation

south of the fault occurred within shallow marine, iatertidal and supratidal environments resullt-

ing from the prevailing. regional regressive-transgressive-regressive marine cycle. Modification

o€ this cycle north of the faulr zone was influenced by contemporaneous faulting.

Introduction

This paper deals with Umberatana Group

sédimentation in a region of contemporaneous

fwulting on the western flank of the Flinders

Ranges, 12 km NNW. of Melrose (Fig, 1),

This is in the southeast quadrant of the

ORROROO 1:250,000 geological sheet map-

ped by Binks (1968). Deposits of the Sturtian

and Elatioa glaciations form the basal and

capping Components, respectively, of the Um-

beralana Group. The intervening period is re-

presenied by a variable sequence of clastic and

carbonate sediments of non-glacigene origin.

{Thomson 1969).

The study area is divided by » fuule zone

into two subareas. of differing stratigraphies,

These reflect pehecontemporaneous movement

on the east-west Spring Creck Mine Fault

(new name). The regional pattern of the urea

was one of a regressive-transgressive-regressive

marine cycle, represented by the stratigraphy

of the southern subarea (Fig. 7}. However,

fault. rupture and sitbsequent subsidence north

of the fault zone produced a short transgres-

sion within the initial regressive phase, but the

main sedimentation cycle regained dominunce

as the sedimentation rate execeded the tate of

subsidence.

Time-stratigraphic relationships, as shown in

Fig. 7, are based on the assumption that the

fault zone cun be interpreted as in Fig. 5. The

nature af outcrop is not sufficent, however, to

fully confirm this interpretation, and the pos-

sibility of complex fault slices of the strati-

graphy contained within the fault zane is not

discounted,

Straligraphic nomenclature (Fig. 2) is that

of Thomson et al, (1964) and Thomson

(1969). Detailed geology is deserihed in

Plummer (1974)!,

*Centre for Precambrian Research, Department of Geology and Mineralogy, University of Adelaide,

Adelaide, S. Anst, 5000.

' Plummer, PLS. (1974). —The siratigraphy. sedimentology und palagoenvironments of Late Precarn-

briatt Uniberatana Group in the Mount Remarkable-Alligator Gorge area, South Australia, Univ,

Adel, linpublished Honours thiesis,

30 P. S&S. PLUMMER & VY, A. GOSTIN

SOUTH AUSTRALIA

Melrose

12 km

j= = = Elatina Formation

fe. sandstones, siltstones

Wilmington Formation

fine sandstones, siltstones, limestones

Angepena Formation

fine sandstones with clay & coarse sand

Brighton Limestone

limestones some silty, dolostones

with clay or coids

Tapley Hill Formation

laminated siltstones

Sturt Tillite

L sandstones, tillites

v\e - \¥ \ \ Md

r\i\ I \ Vy \ eee Fault-rock

‘ a Vite i Wy \W )

Vey A \\y

\ NN Va \V\ Stromatolite

Fig. 1. Generalized geological map of the Umberatana Group in the study area.

FAULTING AND SEDIMENTATION, SOUTHERN FLINDERS RANGES 31

Stratigraphic Nomenclature

Cambrian Elatina

Formation

Wilpena

Wilmington

Group Formation

Angepena

Umberatana Formation

Group Brighton

Limestone

ADELAIDE

Tapley Hill

Formation

Sturt

Tillite

Northern

Subarea

Southern

Subarea

Lithologies

siltstone

fine sandstone

limestone

dojostone

HEGOE

silty limestone

\3/s

Scale in

Metres

silty dolostone

calcareous siltstone

stromatolite

ooid

coarse sand, flat dolomite pebbles

Fig. 2. Stratigraphic nomenclature and general stratigraphic columns of the two subareas divided by

Spring Creek Mine Fault. Thickness variations controlled by subsidence north of the fault.

32 P. S. PLUMMER & Y. A. GOSTIN

Stromatolite biaherm

arbitrary

suriaces

PSP'75

isochrons'

Stromatolite Facies

anes small columnar

BARA (2=5om wide)

large = columnar

ee {7-zocm wide)

SBE stratitorm

25 Intrafermational limestone

Fre. 3. Diagramatic favies sketch of the base and edge of a stramatolite bioherm, Middle Brighton

Limestone, southern subsren.

Sedimentation prior to faulting

Following the Sturtian glaciation, interglacial

sedimentation began in a marine basin helow

the wave base, outside the zone of carbonate

deposition, and under reducing conditions. The

resultant lithologies, namely black, finely lam-

inated pyritic quarizose shales grading upward

into well laminated dark prey quurtzose silt-

stones, form the ‘Tapley Hill Formation

(Howchin 1904; Thomson ef al. 1964).

Gradually the sediment surface rose into the

zone of carbonate deposition, resulting in an

increase in carbonate content toward the top

of this tormation. Eventually carbonate be-

cume dominant as a well laminated dark grey

silty limestone was deposited (basal Brighton

Limestone—Howchin 1904). Above this, the

presence of large dendroid branching stromuto-

lites, set in a blue-grey limestone, testify to

the basin floor rising above the level amenable

to stromutolite growth. Current activity during

this initial period of uniform sedimentation

was minimal, the dominant direction ivitially

being from the north-west, but later swinging

(to Now from the south-west (Fig, 6). A

gradual increase in current intensily is sug-

gested by the change from rare small cross-

laminations (lower Tupley Hill Formation) to

more common shallow ripple marks (basal

Brighton Limestone).

Faulting with sedimentation

Movement ulong the Spring Creek Mine

Fault ended Lower Brighton Limestone sedi-

mentation and produced variations in litha-

Jogies across the fault. A sketch of the fault

zone is shown in Fig. 5. Tentative correlations

of tocks in this zone with adjacent lithologies

suggest two periods of faulting (Fig. 7), the

total stratigraphic throw being approximately

750 m, Fault-zones facies 1 (FFI)—a dark

grey micritic limestone (breeciated in placesi,

with stromutolitic Jimestone megaclasts (Wp to

0.5 m long) of the previously deposited

Brighton Limestone—suggests initial rupture

along, the fault, whereas subsequent facies lack-

ing megaclasts (and FF2 lacking brecciation)

imply warping due to very gradual subsidence,

Seuthern Subarea

South of the fault, shallowing steadily con-

tinucd, At, or just after the time of initial

fuulting, the blue-yrey stromatolitic limestones

of the preceding period grade slowly upward

into massive deep reddish-purple intra-tormu-

tional limestones. containing abundant ‘bull-

yellow stromatolite bioherms, displaying large

furcate to small digitate branching columns, as

the sedimentation surface rose into the zone af

Wave agitation, oxygenation and tidal activity.

Within these bioherms, a zonation based on

stromatolite morphology occurs (Fig. 3), sug-

gesting a series of growth stages tesulling from

energy fluctuations within the depositional cn-

vironment, indicating emergence of the bio-

herms through the intertidal zone in’ the

following manner:

(a) basal stratiform mats lrapped sediment

forming, a stable luyer within the hizgh-

FAULTING AND SEDIMENTATION SOUTHERN FLINDERS RANGES 33

Fig. 4, Tepee structure, Upper Brighton Limestone, northern subarea,

energy subtidal to low intertidal zone;

(b) ‘large’ columns (7-20 em wide) then

developed within the lower to middle

intertidal zone. Abundant carbonate

flakes, originating from disruptive Wave

and tidal action, lodged between these

columns;

bioherms in the high intertidal zone.

Rare disruption of laminae, or an ef-

ficient drainage of debris from the

intercolumn maze explains the relative

lack of carbonate flukes in this zone;

and

(d) stratiform layering on the bioherm

flanks encasing both facies (b) and (c)

enhanced drainage from the centre of

the bioherm,

Fault-zone facies 2, a yellow dolostone,

lacking brecciation and in places sandy, sug-

gesting gradual subsidence and warping of the

still soft sediment. and FF3, a brecciated pale

yellow dolostone, were probably deposited

during this period of stromatolite proliferation.

Continued shallowing then produced a

supratidal environment of deposition and

completed the initial regressive sequence of

the interglacial period south of Spring Creck

Mine Fault. A purple dolostone, with disrupted

clay laminae and lenticular beds of quartz

sand, caps the Brighton Limestone. Fault-zone

facies 4 and 5—brecciated purple dolostones

(with clay in the former)—are the lime

equivalents of this unit, although north of the

fault these two facies possibly represent the

time equivalents of two very different litho-

logics (uppermost Brighton Limestone and

Angepena Formation).

Northern Subarea

At the time of initial rupture along Spring

Creek Mine Fault, the northern block was

downathrown below the depth amenable to

stromatolite growth, yet remaining within the

zone of carbonate deposition, resulting in

laggy grey limestones bearing 4 minor quartz

silt fraction. The thickness (125 m), and uni-

formity of this unit is a result of equivalent

rates of subsidence and sedimentation. Rare

large scale tipple marks (wavelength 50 em.

amplitude 8 cm) and cross-bedding (up to

2.5 m per set with foreset slopes between 15°

and 40°), plus the more abundant small scale

current-ripples and cross-stratification, suggest

an increase in current activity attributed to the

change in seafloor topography. Palacocurrent

analysis reveals several modes us shown in Fig.

6. Rare lenticular outcrops of grey stromato-

litic and purple intraformational limestones

34 P. S. PLUMMER & V. A. GOSTIN

suggest that the depths of oxygenation, agita-

tion and stromatolite growth were almost

juxtaposed.

Above this unit a change in colour and litho-

logy suggests shallowing of the basin to an

oxygenated wave agitated depositional environ-

ment, within which a 20 m thick unit was de-

posited. This unit exhibits facies variation from

pink dolomitic oolites with birdseye structures,

and sandy dolostones near the fault grading

northward into white limestones with dolomite

flakes, generally graded within trough cross-

beds, and rare ooids. A gentle palacoslope

northward away from the fault is therefore

suggested. A supratidal environment followed,

depositing a 5 m thick purple dolostone with

clay laminae exhibiting tepee structures (Fig.

4) in the vicinity of the fault. This unit, pos-

sibly equivalent to fault-zone facies 4 (and ?

5), caps the Brighton Limestone north of

Spring Creek Mine Fault as does its counter-

part to the south. It therefore appears that

supratidal deposition occurred uniformly over

the entire area at this time, suggesting a tem-

porary quiescence of fault activity (Fig. 7).

These purple dolostones were overlain by

350 m of the greyish-red fine sandstones of

the Angepena Formation (Thomson 1969).

These contain abundant purple clay laminae

providing excellent bedding plane partings re-

vealing a wealth of ripple marks and both

polygonal and sinuous mudcracks. Lenses of

coarse-grained rounded sand are common,

often graded and draped with clay. These

sedimentary structures typify a vast tidal flat

to distal fluviatile depositional environment.

The abundance of desiccation cracks, some

reaching 1 cm in width, favours a non-

marine influence. Palaeocurrent analysis re-

veals a dominant easterly drainage pattern with

minor southern and western components (Fig.

6).

This dominantly regressive sequence north

of Spring Creek Mine Fault was then overlain

by the transgressive grey-green deposits of the

Fig. 5. Facies sketch map of the fault zone, lowermost Wilmington Formation (Thomson

Spring Creek Mine Fault. 1969). Siltstone pillows in dolostone are over-

FFI: dark grey micritic limestone with Jain by grey slightly calcareous siltstones with

wtaces megaclasts, brecciated in Chioritic wavy laminae, ripple marks and rare

FF2: yellow dolostone, sandy in places. sinuous mudcracks. These in turn are overlain

by a massive grey calcareous siltstone bearing

FF3: brecciated pale yellow dolostone. ] “

FF4: brecciated purple dolostone with pillow structures, slump beds and _ lenticular

Bal

taal

[NN

—

\=\

clay. beds of olive green-brown oolitic limestone. A

FFS: brecciated purple dolostone. very shallow, yet wholly submerged environ-

FF6: grey silty limestone. ment is therefore suggested. Fault-zone facies

Formation symbols as in Fig. 7. 6, a grey silty limestone, is possibly the equiva-

FAULTING AND SEDIMENTATION, SOUTHERN FLINDERS RANGES 35

-pP=S

a=)

Current Roses

@) Number of readings

Ripple Marks

stippled: symmetrical

Formation

solid: asymmetrical

Cross-stratification

Wilmington

Brighton

PSP'75

Fig. 6. Palaeocurrent analysis of the Umberatana Group in the study area.

Stratigraphy

Southern

Average

Thickness

(mn)

ria. 7,

PS. PLUMMER & Y¥. A. GOSTIN

Fault

Activity

Stratigraphy

Northern

Subarea

downwarping

‘Vime-stratipraphic suramary chart of lithologies, fault activity and palueocenvironments. Lithalo

gies as in Fig, (2. THI: Tapley Hill Formation; BL; Brighton Limestone; AF: Angepena Forma-

tioh; WE: Wilmington Formation; BF: Elatina Formation; L: Lower; M: Middle; U: Upper: dg:

dark grey: g: grey: ert green; r+ red and. purple; w: white and pink: y: yellow.

lent of this period, and represents the final

phase of faulting (Fig. 7).

South of the fault, the Angepena Formation

and the two lower parts of the Wilmington

Formation are absent. The contact between the

Upper Brighton Limestone and the Lower

Wilmington Formation is nowhere visible. It

is suggested that either an hiatus in deposition

occurs between the Upper Brighton Limestone

und Lower Wilmington Formation, or alterna-

tively the Upper Brighton Limestone desposi-

Lier continued in a restricted, slightly elevated

covironment, whilst to the north the much

thicker Afngepenu Formation and the two

lower Units of the Wilmington Formation were

fad down.

Continued warping

Deposition was dominantly ternigenous as

Spring Creek Mine Fault cyolved from a zone

of separation of the two subareas into a flexure

with tore marked subsidence of the northern

subarea, The fault zone was still a major influ-

ence of Wilmington Formation sedimentation

as [ucies changes and thickness Variations uc~

curred above, of immediately adjacent to it.

Massive dark grey very fine sandstones over-

lain hy brown fine and medium sandstones of

the lower Wilmington Formation grade south-

ward into flagey grey siltstones supporting the

palacocurrent data of a west-north-wester'ly

source area. Widespread slumping and cross-

bedding within included sandstone beds imply

a palaeoslape (possibly submarine deltaic)

away from this source region.

The ensuing calcareous member of the up-

permost Wilmingion Formation {herein re-

warded as the equivalent of the Trezona

Formation of Thomson er ai, 1964) repre-

senis the muximum transgressive phase of

deposition. Flagey grey silty limestoncs and

niassive vrey, in places stromatolitic, limestones

in the south thicken rapidly north of the fault

zone (by up to 5 times) and interdigitate with

n buff coarse sandy cross-hedded limestone

bearing ooids and imbricate) dolomite intra-

clasts, The repetitive nature of this interfinger-

ing, which commences directly above the Fault

zone (Plummer 19744, Fig. 10), probably re-

sulled from fluctuating rates of sedimentation

and subsidence within a shallow sea.

FAULTING AND SEDIMENTATION, SOUTHERN FLINDERS RANGES 37

The final regressive coarsening upward se-

quence of laminated siltstones to fine and

coarse sandstones suggest prograding deltaic

sedimentation. Ripple marks, cross-bedding

and some slide-slump bedding support a west

to east palaeoslope, (Fig, 6). Cappimg the

Umberatana Group are the massive purple

siltstones of the Elatina Formation bearing

(?iee-rafted) erratics. Equivalents of these

siltstones elsewhere in the Flinders Ranges are

distinctly glacigene in origin (Elatina glaciation

of Mawson 1949).

Conclusion

Stratigraphic study south of the Spring

Creek Mine Fault reveals that a regressive-

transgressive-regressive cycle of marine sedi-

mentation took place during the deposition of

the interglacial portion of the Late Precamb-

rian Umberatana. Group. The sedimentary

structures and presence of stromatolites within

these sediments indicate that environmental

conditions ranged from shallow subtidal,

through intertidal to supratidal, then back to a

shallow subtidal environment.

To the north of the Spring Creek Mine

Fault, two periods of faulting contemporaneous

with deposition caused water depth fluctuations

during the initial period of shallow subtidal

deposition, then rapid shallowing through

intertidal and supratidal conditions to a mar-

ginal marine (possibly distal floodplain) en-

vironment, This faulting and later subsidence

hinged along the fault zone, caused a 750 m

thickening of the succession to the north of

the fault.

Acknowledgments

The authors wish to thank Dr B. Daily for

critical examination of this text and helpful

discussion in the ficld. Acknowledgment is

also made to ihe Australian Rescarch Grants

Committce for supporting the research of Dr

VY. A, Gostin,

References

Bin«s, P. J. (1968) —ORROROO map shect, Geo-

logical Atlas of South Australia, 1:250,000

series. (Geol. Surv. S. Aust. Adelaide.)

Bryvgs, P. J. (1971).—The Geology of the

ORROROO 1:250,000 map area. Rep. Invest,

geol, Surv',, 8. Aust. 36,

Coats, R. P, (1973.).—Explanatory notes for the

COPLEY 1:250,000 geoiogical map, Geel.

Sty. 5. Aust.

Forges, B, G, (1971),—A table of Adeluidean

and Cambrian stratigraphic names. Quart.

geol. Notes, geal, Surv, 8. Aust. 38.

Howcuin, W. {1904).—The geology of the Mount

Lofty Ranges, Part 1. The coastal district.

Trans. R, Soe. §, Aust. 28, 253-280,

Howcuin, W, (1916).—The geology of Mount

Borariabie: Trans. R. Sac. §. Ausi. 40, 545-

Mawson, D. (1949).—Elatina glaciation: a third

recurrence of glaciation evidenced in the

Adelaide System. Trans. R. Soc. S. Aust. 73,

117-121.

Smuru, D, B. (1974) —Origin of tepees in Upper

Permian Shelf carbonate rocks of Guadalupe

Mountains, New Mexico, Bull. Am. Ass. Per

rol, Geal. 58, 63-70.

Tuomson, B. P. (1969)—Precambrian Basement

Cover—The Adelaide System. Jn L, W., Par-

kin {Ed.), “Handbook of South Australian

Geology”, pp. 49-83. (Geol. Surv. S. Aust.:

Adelaide).

TuHomson, B. P., Coats, R. P.. MirAms, R. C.,

Forses, B, G., DeiGarno, C, R., & JOHNSON,

J. BE. (1964).—Precambrian rock groups in

the Adelaide Geosyncline: A new subdivision,

Quart, zeal. Notes, geol, Surv. 8, Aust, 9.

SMALL FOSSIL VERTEBRATES FROM VICTORIA CAVE,

NARACOORTE, SOUTH AUSTRALIA

IV. REPTILES

BY MEREDITH J. SMITH

Summary

SMITH, Meredith J. (1976) .-Small fossil vertebrates from Victoria Cave, Naracoorte, South

Australia. Trans. R. Soc. S. Aust. 100(1), 39-5 1, 28 February, 1976.

Reptile fossils have been found at Naracoorte, South Australia, in a Pleistocene cave deposit that is

rich in marsupial and rodent remains. Reptile vertebrae are abundant and a few jaws and limb bones

have been recovered. The diagnostic features of these bones are described.

Of the twelve reptile species present, nine still live in the Naracoorte area; they are three elapid

snakes, Pseudonaja c.f. P. nuchalis, Notechis c.f. N. scutatus, and Pseudechis c.f. P. porphyriacus;

and six lizards, Varanus varius, V. gouldii, Trachydosaurus rugosus, Tiliqua nigrolutea, Egernia

c.f. E. whitei and a species consistent with Sphenomorphus tympanum. A fourth elapid snake

represented by 40 isolated vertebrae, and a species of Amphibolurus have not

been identified.

The remaining species is a boid snake, described here as a new genus and species, Wonambi

naracoortensis. The eight vertebrae recovered are large, suggesting a length of at least 5 m for the

whole snake. Morphologically, the vertebrae differ strongly from those of other Australian boids in

having a high but back-sloping neural spine, paracotylar foramina present, accessory processes

absent, and, particularly, in having large paradiapophyses that extend further laterally than the

zygapophyses. These vertebrae closely resemble those of Madstoia bai Simpson from the Eocene of

Patagonia, but without cranial remains of both species, no relationship can be postulated between

Wonambi and Madstoia.

SMALL FOSSIL VERTEBRATES FROM VICTORIA CAVE,

NARACOORTE, SOUTH AUSTRALIA

Ty. REPTILES

by Merepiru J. SMITH

Summary

Smiry, Meredith J, (1976),—Small fossil vertebrates from Victoria Cave, Naracoorte, Sowih

Australia. Trans. R. Sec. S. Aust. 100(1), 39-51, 28 February, 1976.

Reptile fossils have been found .at Naracoorte, South Australia, in a Pleistocenc cave

deposit that is rich in marsupial and rodent remains. Reptile veriehbrac are abundant and a

few jaws and limb bones have been recovered. The diagnostic features of these bones are

described.

OF the iwelve reptile species present, nine still live in the Naracoorte area; they are three

tlapid snakes, Psexdonaja c.f. P. nuchalis, Notechis c.f. N. scutatus, and Pseudechis cf. P. por-

phyriacus; and six lizards, Varanis varius, V. gouldii, Trachydosaurus rugosus, Tiliqua niero-

lutea, Egernia ct. E, whitei and a species consistent with Sphenomorphus tympanum. A fourth

elapid snake represented by 40 isolated vertebrac, and a species of Amphibolurus haye not

been identified.

The remaining, species is a boid snake, described here as a new genus and species,

Wonumbhi naracoertensis. The eight vertebrae recovered are large, suggesting a length of al

least 5 m for the whole snake. Morphologically, the vertebrae differ strongly from those of

other Ausiralian botds in having a high but back-sloping neural spine, paracotylar foramina

present, accessory processes absent, and, particularly, in having large parudiapophyses that

extend further laterally than the zygapophyses.. These vertebrae closely resemble those of

Madstoiq bai Simpson from the Eocene of Patagonia, but without cranial remains of both

Species, no relationship can be postulated between Wonambi and Madstoia,

Introduction

The reptile faunas of Australian fossil

deposits have rarely becn completely anatysed.

For some deposits, the presence of unidentified

repliles has been noted (c.g, Archer 1974,

Dortch & Merrilees 1971, Gill & Banks. 1956,

Lundelius 1963); for other deposits, the most

distinctive species have been identified, but

often ta genus only (eg, Merrilecs 1968;

Thorne 1971). Exceptions are the carpet snake,

Python variegatus (=Python spilotus varie-

gets) associated with the extinct marsupial,

Thylacoleo sp., and other marsupial remains at

Marmeor Quarry, Queensland (Longman 1925)

and the sleepy lizard, Trachydosaurus rugosus,

at Gore Limestone Quarries, Queensland

(Longman 1945), Remains of a large extinct

Varanid lizard, Megalania prisca, have been

found in Pleistocene deposits in Queensland,

New South Wales and central Australia (Fejer-

vary 1918, 1935; Hecht L975).

The deposit in Victoria Cave at Naracoorte,

South Australia, is probably of Pleistocene age

(Smith 1971). Among the large animals,

eXtinet species are common (yan Tets & Smith

1974; Wells, pers. comm.), but, in contrast, the

smal] marsupials and small birds are referrable

to modern species, though not all of them

occur in the Naracoorte area now (Smith

1971, 1972; van Tets & Smith 1974),

Por identifying reptile species, characteristics

of skull fragments, jaws and teeth are of less

value than they are for mammiails. As reptiles

grow throughout life and are polyphyodont, the

“adult” dention cannot be defined us it can in

mammals. The variations in dentition between

species in many genera is no greater than

within species. Fortunately the vertebrae of

reptiles are of diagnostic value and Auffenberg

(1963) was able to identify single vertebrae of

North American snakes to genus and often to

species. Diagnostic vertebyal characters have

* Department of Zovlogy, University of Adelaide, Adelaide, SA. 5000.

40 MEREDITH J. SMITH

accessory

process colyle

subcentral foramen

prezygapophysis paratiapophysis

hypapophysis: postzyyapophysis

zyyantrum—_ eandvla

zyausphen’ neural spine

Jateral foramen

arapophysial

oA hall centrum

cundyle

Elapid Varanid Scincid Agamid

Fig. I. Ventral (above) and lateral views of precaudal vertebrae of (a) Pseudonaja nuchalis

(Blapidae), (b) Veranus gouldit (Varanidae), (¢) Tiliqua occipitalis (Scincidae), (d) Amphi-

bolurus barbatus (Agamidae).

Fig. 3. Vertebrae of Wonambi naracoortensis arc distinguished by their wide paradiapophyses (p) and

presence of paracotylar foramina (f) as seen in (a) anterior view of P16144k and (b) lateral,

and (c) posterior view of P16i44s.

Fig. 5. The prezygapophysial facets are less upturned in Trachkydosaurus rugosus (6) than in Tiliqua

nigrolutea (a).

Fig, 6. Distally, the fused sacral pleurapophyses are cup-shaped in Trachydesauras rugosus (a) and T,

occipitalis (b) but are triangular in T. nigralutea (c), T. scincoides (d) and Egernia cunnine-

hami (€).

not been established precisely for any Aus- Methods

tralian snake species (Smith 1975). Fossil re-

mains of lizards include not only skull bones

and yertebrae but also some limb bones and follow, Smith, (1971).

Methods of collection and preservation

some elements of pectoral and pelvic girdles, The present maximum depth of excavation

These are briefly described, and their diagnos- is 80 cm, although hone chips occur in cores

tic values assessed in this paper. taken as deep as 2,5 m,

SMALL FOSSIL REPTILES FROM NARACOORTE 41

TABLE 1

Some dimensions of the holotype (P16168) and the seven paratype vertebrae of Wonambi naracoortensis

(Dimensions in mm)

Specimen

P16168 P16170a P16166 P16129t P1l6144k P16144s

P16167 P16170b

Length between zygapophyses 16.5 15.4

Height (centrum + neural spine) 32.3 34.1*

Width across prezygapophyses 25.6 21.1

Width across paradiapophyses 28.5 22.3

Minimum width of centrum 18.6 15.9

Width of zygosphene 9.2 8.4

Width of condyle 10.8 8.2

Length of prezygapophysis 8.1 6.1

* Height includes length of hypapophysis.

Skull and jaw elements of all but the most

robust species were rarely recovered from the

Victoria Cave deposit, whereas vertebrae were

common. Consequently, for the diagnosis of

reptile species in this fauna, vertebrae have

been considered in detail and other bones more

briefly. Comparisons have been made with dry,

disarticulated skeletons, and occasionally with

cleared, alizarin-stained whole specimens.

Descriptive terms (Fig. 1) follow Auffen-

berg (1963). The “length” referred to in des-

criptions of vertebrae is the greatest distance

from the anterior edge of the prezygapophysis

to the posterior edge of the postzygapophysis

(Pr-Po of Smith 1975). The ranges of lengths

are given, with mean and standard error.

Measurements were made to the nearest 0.1

mm, with dial-reading, needle-point calipers.

The fossil specimens are lodged in the South

Australian Museum (SAM).

Results

Family BOIDAE

Boid vertebrae lack parapophysial processes,

the accessory processes are very short or absent

and the vertebrae lack hypapophyses on the

posterior two thirds of the precloacal column

(Hoffstetter & Gase 1969).

Wonambi n. gen.

Definition: Vertebrae characterized by a high,

backwardly-sloping neural spine; slightly up-

turned zygapophysial facets; large paradiapo-

physes extending laterally beyond the zygapo-

physes; and a pair of paracotylar foramina.

Type species: Wonambi naracoortensis

Content: W. naracoortensis is the only known

species in the genus.

“Wonambi” is derived from an aboriginal

name for the mythical rainbow serpent (Elkin

1964).

18.1 16.4 19.3 21.8 23.4 19.6

40,8* 33.1 38.4 37.1 41.0 36,2

27.7 25.8 30.4 33.0 35.5 29.5

29.7 27.7 33,2 41,1 43.7 33.0

21.2 19.1 22.7 24.1 27.1 21.8

11.8 9.3 12.6 10.9 12.4 11.3

10.6 10.3 12.6 12.3 12.7 12.4

1.5 74 8.9 11.8 12.8 10.9

Wonambi naracoortensis n. sp.

Holotype: SAM, P16168. A dorsal vertebra

collected in Fossil Chamber, Victoria Cave,

Naracoorte, S. Aust., at a depth not greater

than 30 cm below the surface of the cave

earth.

Definition: The same as for the genus Wonambi

until other species are described.

Description: The neural spine is high (Fig.

2C); its anterior edge begins near the rim of

the zygosphene and rises obliquely to the hori-

zontal dorsal edge of the spine. The spine over-

hangs slightly posteriorly. The zygosphene is

narrow (Table 1) but it is so heavily thickened

that it is as deep in dorsoventral extent as it is

broad. The zygosphenal facets are almost ver-

tical (c. 70° to horizontal) (Fig. 2D). The

relatively small zygapophyses are slightly up-

turned (c. 25° from horizontal). Accessory

processes are completely absent. The paradia-

pophyses are large (Table 1); the upper part

of the articular facet is convex and protrudes so

far from the centrum that the maximum width

of the vertebra is the width measured across

the paradiapophyses; the lower part of the

articular facet is flat. The cotyle and condyle

are slightly depressed (Fig. 2A) and the top of

the condyle is tilted forwards at c. 75° to the

vertical. The ventral surface of the centrum is

smoothly rounded, with weak subcentral ridges

and a low median ridge that terminates pos-

teriorly as a blunt haemal keel, notched in the

midline (Fig. 2B).

Each foramen of the subcentral pair is

located close to the median ridge at about mid-

centrum; each foramen of the lateral pair lies

on the neural arch pillar about halfway between

paradiapophysis and postzygapophysis. There

appear to be two pairs of parazygantral fora-

mina, but, as the bone is pitted in this region.

42 MEREDITH J. SMITH

Fig. 2. The holotype of Wonambi naracoortensis (P16168) in dorsal (A), ventral (B). lateral (C), an-

terior (D), and posterior (E) views.

SMALL FOSSIL REPTILES FROM NARACOORTE AR

foraming are distinginshed with difficulty fron

pits. ‘lhe two foramina of a paracolylar pair

lie close to the rim of the coryle near the top

of the centrum,

Fariation; The paralype series consists of seven

dorsal vertebrac. P16129t, Pl6i44k,s, P16166-

7, and P16070a, b all but une of which were

found in the top 30 cm of the cave carth of

Fossil Chamber, Victoria Cave. All specimens

share with the holotype the diagnostic features,

especially ihe widely spaced paradiapophyses

(Table 1, Fig. 3), The parazygantral foramina

consist Of a single pai which are sunk into

hollows in P16129s, P16167 and P16170b; and

m Pl6144k the single pair of zygantral fora-

mina can be seen clearly. The haemal kcel is

notched posteriorly im P16129s, P16167 and

PL6170b (as in the holotype), these vertebrae

having occurred nearer the tail than the others,

ws judged from their lower neural spines.

PI6166 bears a small hypapophysis. and

P16170a a longer hypapophysis.

Associated material: A fragment (P16170c)

from near the anterior end of a Teft maxilla

with three teeth, curyed backwards, Each tooth

is approximately 7 mm long.

Assuming thal PL6I67 represented tlre

largest vertebra of the specimen of H*onaebi

raracoortensis, and assuming that the largest

vertebrae of HW. narieooriensis and Python

sPilotus, respectively, occupy the same propor

tion of the length of the vertches! cofomn, then

PIA167 would have been derived from 3 snake

of total Jength about 5.0 m,

Comparison with arher species; OF the cight

Australian species of Boidae., vertebrae of

Python spilomes (4 spovimens), P, ametiustinuy

(2), biavis childrent (1), Chondropy ten

wirtdis (1) and Aspidites melanocerhalis (3)

have heen examined,

The vertebrae of these extant hoids are

vharacterized by the presence of small. pointed,

avcessiry processes beneath the prezygapo-

physes, by having large outwardly-directed

zygapophyses exteniling Further laterally than

the pecadiapophyses (Table 2) and the neural

spine hatchet-shaped. and hy the absence of

paracotylar foramina, Subgentral ridges are

strongly developed. The general shape of the

vertebra is similar among all the spovics

(Tahles 2, 3). Chandrepython viridis diflers im

having the neural spine bifurcate anteriorly,

Pyrhou spilotus aid P. untethistirins resemble

euch other in having a pair of foramina at the

base of the neural spinc, while Asplidites

nelarocephalits, Chondropython virtdiy and

Liesiy ofildreni Yack thls pair of foramina.

Wonamhit natacoertensis differs fram all

other Australian boids in Jacking accessory pro-

cesses, iu having the neural spine sloping buck-

wards, in having weak subcentral ridges, and

in the presence of paracotylar foramina, Jt

shares with spiditey melanacephalus. Chen-

dropython viridis and Liasis ofildrent the

absence of foramina at the base of the neural

spine, The totul height (relative to length) of

the vertebrae is greater in HW. naracoartensiz

thin in other Australian boids, the paradiapo-

physes extend further lateratly than the zygapo-

physes, the condyle (relative to Vertebra

length) is wider and the zygosphene (relative

to vertebra length) is narrower (Table 2). On

the other hand. width across prezygapophyses,

Minimum width of centrum, and length of pre-

zyezapophysis (all relative to vertebra length)

Fall within the ranges of the extant species, as

does height/ width of condyle (Table 3).

Paracotylar foramina wre generally absent in

extant boids, and occur only in the genera Con-

stricror and Trepidophls (Boinac) and Pnygrus

(Erycinael (Hoffstetter & Guyrard 1964).

They are found, wsually as two pairs, in the

fossil genera Giyantephix and Madstoia (Hoff-

steller 1961a, hb}, and as a single pair in HW,

NEPUCOOrLIPASIS.

Whereas the lengths of vertebrae of

Wonornhé iaravcortensis (relative to width

across prezVgapophyses) fall within the ranges

of those of the extant Australian boids, inelud-

ing Liasis (Table 3), six vertebrae from the

Weillifigion caves of New South Wales were

langer than vertchrae of Liaws (tyidekker

1888, p. 256).

There is a striking resemblance hetween

Honaithi vertebrae and those of Mudsreia tial

(Palaencene-Eocenc of Palagenia) and M,

madagascariensis (Cretaceous. Madagascar)

(Hofstetter 1961a, Simpson 1933), particu.

Jarly in the back-sloping neural spine, broad

paradiapophyses and absence of accessory

processes, Hoffstetter (1961la, b) included

Madsraia and Gigantephis (from the Eocene

of Egypr, Andrews 1906) in a sub-family Mad-

stoiinac. The diagnostic features were: (a)

accessory processes absent; (b) a pair of para

zygantral foramina present and opening into

deep hollows; and (c) paracotylar foramina

always present, usually as two pairs. The two

genera were distinguished by the form of the

ventral surface of the centrum, (ffeantephir

having an undivided haemal keel, and Afad-

44 MEREDITH J. SMITH

sioja having the haemal keel distinctly divided

into two. All WY. waracoorlensis vertebrae con-

form with character {a) and differ from «ther

hoids, all of which have accessory processes

(Hoffstetter 1961a); and the more posterior

vertebrae, conform with (bh). However all the

vertebrae have distinct parazyyantra) foramina

(even though they emerge through the charue-

teristic deep hollow only in the posterior verte-

frac) and such foramina in modern boids arc

minute, irregular and inconstant (Hoffstetter

61a). W. naraceorteisis vertchrac differ

slightly from (c) in having « single pair of

paracotylar foramina, but their presence at all

ig rare in boids (Hoflstetter & Gayrard 1964),

The form of the yentral surface of W. nara-

coortensis vertebrae ranges from hypapophysis

prescot {P16166) or haemal keel undivided

(P161291, P1éid4k}, to haemal keel notehed

(P16168) and finally to haemal keel distinctly

divided (P16129s and P16167); thus it encom-

passes the form of both Muadstefa and Gigan-

tophis. Medstoia differs fram Gigantaphiy also

in the greater development of neural spines and

Parsdiapophyses (Simpson 1933). In these

features WB’, naraceertensts closely resembles

Mad sraie.

When Madstoia was compared with many

boids, both recent and fossil, the resemblance

af Madstoia to Gigantophis was found to be

closer than to other known genera. However

it was impossible to conclude that the two were

definitely more closely related to each other

than to other fossil boids (Simpson 1933),

Similarly the relationship of (Wanarmbi to Mad-

stoia or any other boid will remiuin obscure

wml the skull is known.

The presence of Madstoie in Patagonia and

Madagascar has been regarded us evidence of

former continuity of the southern continents

(Hershkoyitz 1972, p. 316).

Family ELAPIDAE

Elapid vertebrae have conspicuous accessory

processes and hypapophyses on all precloacals

(Fig, 1).

Psendonaja c.£. P. nuchalis Guother

Material: Vertebrae (566 precloacal, 25

cuudal); dentaries (6),

The vertebrae have heen described (Smith

1975). The largest with a teneth of 11.1) min

between zygapaphyses would hnve heen derived

frnm a snake about 190 cm long,

The dentary of P. wuchalis is almost stratcht

posteriorly. but anteriorly it curves outwards

then inflects sharply. The teeth are strong and

curve backwards very slightly. They are

separated hy w distance equal to c, 2/3 of the

adjacent teeth, The second tooth is the lonuest

but the succeeding teeth along the dentary de-

crease in size anly slightly. The fossil dentaries

are similar.

Notechis c.f, N, seutatus (Peters)

Material; Vertebrac—precloucal (15,

6.3-9.9 mm, mean 7.6 + 0.35).

These vertebrae differ from P. auehalls and

resemble N. sertates in having a relatively short

neural spine overhanging both anteriorly and

posteriorly (Smith 1975).

length

Pseudechis c.f. P, porphyriacus (Shaw |

Material: Vertebrae—preclouval (55, length

4.1-9.5 mm, mean 6.3 + 0.15); maxillae (2);

dentary (1),

Both Notechis and Prendechis have 3-5

Small teeth, whereas Pseuwdonaje has 8-10

(Worrell 1963; pers, ohserv.). A left maxilla

(Pl6é164a) bears a curved fang, followed after

a diustema, by three small, curved teelh and

is consistent with P. porphyrivueus in size an

shape. A ymaller fragment of a right maxilla

(P16164b) is probably from the same skull,

having been taken from the same sample.

P. porphyriaeus dentaries differ fron P.

nuchalis and N. seutatus dentaries in being

more sharply curved anteriorly; the teeth arc

fine, backwards-curving and closely-set. A right

dentary (P16132c) conforms with FL perphy

raeas,

The vertebrae have the long, acute, accessory

processes (Smith 1975) typical af P. por

riacus but they differ in having these processes

directed more anterolaterally than in the P

porphyriacas available for comparison,

Undetermined

An unidentified elapid group contams 40

vericbrae (length 3,8-7,8 mm. mean 60 +

-'5) characterized by the short, blunt hypapn-

physis.

Family VARANIDAE

Varanid vertebrae are distinguished by the

overhanging condyle (Fig. 1),

Varanus varius (Shaw)

Material; Vertebrac—cervical (4, lengths 18.7-

26,3 mm. mean 21.35 + 1.76), dorsal (17.

lengths—Table 4), sacral (3), caudal (26,

lengths 7,9-19,2 mm, mean 12.3 + .53);

maxilla (19; dentaries (5); pametal (1),

The dorsal vertebrae of Varanus elganrtenus

ane readily distinguished by their broad cencrit

SMALL FOSSIL REPTILES FROM NARACOORTE 45

[ratio Of width across prezygapophyses (Pr—Pr)

to Tamimum width of centrum (BW) < 1.6

(Table 4)] and jong neural spines, vertical both

anteriorly and posteriorly. But the vertebrae of

i. varius and ¥ gonldi are similar morpho-

logically—-there is overlap iv the relative width

of centrum, relative width of candyle (CW)

and telative width across prezygapophyses fall

relative to the length, Pr—Po) and also in the

ratio of width across prezygapophyses to rmini-

mum centrum width (Table 4), These values

overlap even when the comparisan is made

hetween vertebrae fram the same position in

the column The neural canal. viewed from the

front, is slightly depressed in ¥, gonldié bul is

round in . varius.

The fossil dorsal vertebrae are consrstent

with hoth V_ vartuy.and V. eouldil in their pro-

proportions and have the neural canal round

anteriorly, as in VL peiriier

Like the dorsals, the cervical and catidol

vertebrae of VL verins and ¥. gould{i are almost

(Or quite) indistinguishable as ta species, but

the first sacral vertebrae are distinctive. The

iransverse processes of the first sacral of V-

souldi (2 specimens examined) bear several

low. ridges— one such ridge from the anterior-

mose point of the lateral surface of the trans-

verse process extends towards the cotyle: a

diagonal ridge passes from the prezygapophysis

to the lateral postero-darsal tip of the trana-

verse process and a ridge from the Iaternl

altere ventral tip of the transverse process 19

the condyle makes the posterior surface of the

transverse process slightly Cancave, In contrast.

the transverse processes are smoothly-rounded

and convex in VF. varius (3 specimens

examined), Similarly in the fossils {P16135r,

Plé169a) the transverse processes are

smoothly rounded, The fossil conforms with

Vo varius and differs from WV. gouldii also in

having the neural canal round anteriorly and

the transverse processes at their lateral extremi-

tres flared to helow the Jevel of the centrum

(whereas In V. goalddit the flaring extends more

ilorsally }-

In Parenus varus the parictal foramen lies

in the middle third of the length of che parietal

plite (Mertens (942), whereas in FV geulaii

and V. vigdntéus it is in the antetior third

{pers., observ},

Laterally compressed, recurved, pleurocdont

leeth with striated bases are characteristic of

varanids {Edfiund 1969). The teeth of PF,

siemens are fine and thin, but in B. varias and

VF pouldii, and in the fossils, the lateral com-

pression ig less extreme, and a labial and a

lingual ridge ascend each tooth, The hasal

fluting extends nboaut 1/35 of the way wp [he

tooth.

The length of the larger fossil lirst sacral

vertebra (18.8 mm Pr-Po) indicates a total

length of c, 1,6 m for the ammal,

Varanus gouldii (Gray)

Material) Humerus (1).

The shaft of the humerus is smoothly

rounded in P. verits (2 specimens) byt in

V. gouldii (3 Specimens) 2 distinct ridge

extends from the proximal termination of the

supinaive crest to a muscle scar (presumably

for the humeraradialis muselé) near the proxi-

mal expansion, Anteroventrally, he deitopec-

toral crest ig promiment in both V, varivs and

V. garldit, but in ¥. 2gouldti the erest extends

further proximally than in ¥. varins, The fossil

hiimerus (P16146b) conforms with V. gouldél

and dilfers (rom V, varius in having a ridge

extending proximally from the stpinator crest

and apparently also in the proximal develon-

inent of the deltopectoral crest, although most

of the proximal articular facet of the fossil

humerus has been tost. The fossil hay a distinct

tuhercle at the proximal termination of the

supinulor crest. Stich a distinet tubercle was

seen only inane modern specimen of Varanns

species, viz. a very large VL gauldii. No

lubercle could be distinguished in two

pouldii comparable in size with the fossil, nor

in iwo F, verizis.

Family SCINCIDAE

In scincid vertebrae the centrum tapers, in

ventral outline, frorn broader aliteriorly to

narrower posteriorly, and there is no precandy-

lar constriction. The ventral surface of the cen

trum ts smoothly rounded (Fig, 1),

Trachydosanrus rugosns (Gray)

Material: Osteoderms (several hundred); verte-

bra—cervical (10, lengths 5.2 7.0 mm, mean

6.0 + 4.21), dorsal (46, lengths 6.1-10,7 mim,

mean 8.9 — 0.17), sacral (5 pairs), pygal (A.

lengths 6.7-10.0 mm, mean 8.0 + 0,53),

caudal (5, leagths 5.0-7.0 mm, mean 6.3 =

0.373; maxillae (5): premanillac (2); dentaries

(6): humeri (3), femur (1); frontals (24.

Oxteoderms: In Trachydosaurus, the astee

derms are thick und coarsely pitted, whereas in

Tiliqua migrolurea, TU. oevipitalix ynd T. xcin-

coaldes the osteodernms are thinner and finely

pitied; in Everaia cemminghami the darsal asteo-

derms bear a posterior median tooth; and in

4G MEREDITH J. SMITH

Frequency

| Fossil

T. nigralutea (3)

f T. scincoides (3)

T.occipitalis <2)

Fig. 4. Frequency distributions of some dimensions of dorsal vertebrae in which Trachydosaurus

rugosus differs from Tiliqua specics. The sample of fossil vertebrae assigned to T. rugosus has

frequency distributions similar to the modern sample. (a) width across prezygapophyses divided

by length between zygapophyses, (b) width across prezygapophyses divided by maximum width

across paradiapophyses, (c) width of condyle divided by width across postzygapophyses, (d)

height of condyle divided by width of condyle. Although the number of presacral vertebrae with-

out a hypapophysis is 30-32, not all vertebrae could be measured in every specimen.

most other skinks the osteoderms are thin and

almost smooth, except over the head of some.

Cervical vertebrae: Cervical vertebrae of scin-

cids have the hypapophysis sutured or fused to

the posterior part of the centrum whereas in

agumid cervical vertebrae the hypapophyses

are sutured or fused to the anterior part of the

centrum (Hoffstetter & Gase 1969). Cervicals

of Trachydesaurus rugosus have broad,

roundish zygapophysial facets whereas in

Tiliqua species and Egernia species the zygapo-

physes are usually anteroposteriorly elongated

and narrow.

Dorsal vertebrae: Cervical vertebrae are

defined as those anterior to the first vertebra

of which the rib joins the sternum (Hoffstetter

& Gasc 1969), but because this distinction can-

not be applied to isolated vertebrac, I have

included in the discussion of dorsal vertebrae

all the presacral vertebrae that do not bear a

hypapophysis. Dorsal vertebrae of T. rugosus

are squarish in dorsal outline, i.e. the width

SMALL FOSSEIL REPTILES FROM NARACOORTE Aa

across the zygapophyses approximately etuals

the length between the zygapophyses, whereas

dorsals of Tili¢a species are longer than wide

(Fig, 4a), In other skinks also, und even in the

largest Australiag skink, the heavily-built

Evernia buagana {only one = specimen

examined), the vertebrae are longer than wide.

In T. regosuy vertebrae, the neural spine is

low, slopes backwards and overhangs pos-

teriorly, At its posterior termination, the neural

spine is thickened and marked by a short

median groove. The pre-zygapophysial facets

are directed dorsally at an angle to the hori-

zontal of abotit 20-40° (although in the an-

terior 3 or 4 dorsals the angle may be as great

as 60" in some specimens (Fig. 51}. The pre-

zyzapophysial facets extend laterally nearly as

fur as, or further than, the paradjiapophysial

convexities (Fig. 45). The zygapophysial facets

are almost round im contrast with the out-

wardly directed zygapophyses of agamids, The

condyle is narrow [Fig. dc) and slightly

depressed (Fig, 4d). Except for the neural

spine and slight zygapophysial ndges, the cen-

trum is smoothly rounded, again in sharp con-

trast with agamids where not only are the

lateral ridges strong, but also a wide micdventral

ridge is conspicuous (Fig. 4d). The ventral sur-

face of the centrum is triangular in T. rugosyy

(as in agamids} whereas in other skinks, ven-

trally the sides of the centrum are almost

parallel behind the paradiapophyses.

Sorral vertebrae; The pleurapophyses. of the

Sacral vertebrae are fused distaliv for about onc

third of their Jength. The facet for articulation

with the ilium is copped and differs from the

triangular facets of Tiligna species (except T-

orvipiteliy) and Eyernia species (Fig. 6). Tn

T. arcipitalis where the lateral articulation

cupped (as in TL rngoses), the zygapophysial

facets are sharply upturned, at an angle of about

JAQ-S(I", as in other Tiligua species, wherens in

T, rugosus the facets ure only slightly uptuned

(angle c. 20°). The condyle is narrower in T.

rugester than in Tiligna species,

Ceudal vertehrae: T, rugesns eaudals are robus:

und the transverse processes project only

slightly venically, much less venirally than in

T. selecolder, 1. aceipitalis and T. nigrolatec.

Teeth and teoth-bearing bores; The tecth of

Trackydoxeuris resemble these of Tilique

species in having conical tips, whereas tecth of

Eserniat species are laterally compressed at ihe

tips (Mitchell 1950), Usually the teech of 7.

ravosuy are broad atd bunt, but in some

specimens the teeth are longer, Itinner and

sharper, These latter overlap m form With those

of the larger Tiligua species. In T. gerrarai

one tooth in each jaw is very large, about four

Himes the thickness of the others, which tre

fine, with rounded tips. The maxillary bone of

T. rugosux is robust. Beneath the orbit, a strang

bone ridge ruins parallel ta the jaw margin and

extends posteriorly beyond the level of the end

of the tooth row. In Tilique species, this ridge

is) Weaker and shorter, not extending beyond

the end of the tooth row. and often ending still

more anteriorly. The demiaties. toa, are rohust.

and are thicker and deeper, especially an-

teriorly, hear the symphysis, than are those of

Tilique species.

Frontals; Two frontal bones were cach charac-

terized us of T, tuegosus by the (ick, coarsely-

pitled osteoderm fused with the bane,

Limh hones: Homerus aud femor have rela-

tively thick shafts in J) egos,

Tiliqua nigrelotes (Quoy & Gannard)

Material: Veriehrae—dorsal (6, lengths 6.1-

8.9 mm, mean 7.8 + 0.44), pygal (2, length:

5.0, 6.2 mm), caudal (3, lenaths 5.6, 64, 68

mm}; maxillac (4); dentares (3): parietal

bone (13,

The size of the fossil dursal vertebrae indi-

cafes that they were derived from a lizard at

least 22 cm in snout-vent length, Such size is

reached by the larger species of Eyernia and

Tiligva, but not by TT. caruerinae, T-

branchiale, T. peterst nor T. woorl-fonesh.

In Tillqua species, the prezygapophyses are

dozsally upturned at an angle of 35—55° (Fig.

5n) of even greater in the first two or three

dorsals. The dorsal vertebras of Australian

skinks, other than Trachyderavrns, are longer

than wide, except somelimes the Last presacral

which may be slightly wider than long {Fig

4a). In T. seinceides. most of the dorsal verte-

brac (except for the first two or three and last

two or three) are extremely elongated (Fix.

4a) and this species is further characterized hv

the broad, depressed condyle (Fig. 4c, 4) and

narrow zygapophyses directed almost anterod-

posteriorly. In TJ. seineoides, the paradia-

pophyses extend laterally well beyond the

lateral edges of the zvgapophyses (Fil, 4b), In

T. nigralytea und T. oceipitalix the 2zy¥yapo-

phystal facets are slightly wider (though the

width never equals oor exceeds the length) and

srt directed antero- or postero-lalerally, henve

the width across the zygspophyses is greater

(relative to, eg. the length between zvgapo-

MEREDITH J. SMITH

TABLE 2

Vertebral proportions of Wonambi naracoortensis competed with thuse of Jur extant -boids, Ten. verithrak were measurad from the posi-hypapaphystat region of madere

Specimens, The range is followed 2y mean + standard error in pureniheris.

Width across:

paradiapophysés/ Width

Specimen Length (mm) Total height /length across pre-2¥gapophyses Widti condyletLength

WY. naracoortensts

P16168 16,5 1.95 it O45

PL6l4si 194 wy 1.09 0.65

P6144» “26 4,70 124 056

F161294 1h 2,02 1,07 0.83

Pi6167 ma 4,76 7a 054

P1617b we 1,85 142 0,63

Liashs children 43=5,2 (4.9 + 09) 101-433 (1.47 03) 0.90-0:91 10.95 = OL) 364.52 (O47 + 02)

Python spileliis

B T5- 85 (6th AN) HOmeL TY 17) 0.777 0.79 + A) DATHLSS (0.50) + 01)

2 $1- 8.5 (84 08) 124-047 (1.46 le 02) O,7841,40. (0,79 002) O.SIALS8 (AG + oO)

3 11.4-42,0 (11,6 — 07) 1,53-1.-71 (1.61 02) 0.70-0,73 (0.73 O12) 0.474151 (0.49 + 003)

Prrhow amethiccinas

1 145-155 (14.9 =p fa) 132-145 (LaR + OLY D.75-0,78 (U.7T J: 003) 0.47-40.51 (0.49 + (4)

o* 10.1-10,5 (10,3 + 04) 142-249 (1.45 + On) O794),82 10.90 + 009) 0:59-0.62 (0.60 + 003)

Aspidites melancvcephalus

Chondraphysican viridis

oe ————_— eae

* Spécimen incomplete: only aboul one quurter of the precloacs! yertetrac were studied,

37-65 (63 077

35-64 (64— 10)

554 69 (HA td)

139-154 (1.46 4-02)

129-15 (1.43 04)

0.95-1,a5 (1,22 + ,04)

OQ. 7S40.84 (192 + AMG)

0.30-D.86 (089 + 01)

0.73-0,83 (0,78 = 01)

TABLE 3

Vertebral proportions of Wonambj maracanrtensig compared with those of four exsant buids, Ver vertebrae were measured from the post-kypapophysial repian of maders

specimens, The range iz jotiawed by mean + siandard error im parenihesis,

{LSSH1b2 (UY LY

D.S4).62 (0.58 + 01)

O4AD (0.46 — 01D

Zygosphene width)

Length

0.55

0.65

0.50

O07

0.53

Onn

O.S9474 TLAB + ANZ)

0,57-0,72 (0.66 + 02)

{1.65-D.76 (0,71 = OL)

0.7841,90. (0,45 + 01)

O H8-U 72 (0,69 + 14)

O.AS-41.85 (0.68 SE M2)

0,040,77 (0,75 + 01)

USK AGT (G2 HD)

0.5260 10,59 + 01)

W. across pre- Helght condyte/ L. pre-

Specimen zygapiphyses/Length Min w. centram/ Length Width condyle zyRapophysis/ Length

W, naraccortensis

PIBL6S 1s Ln 0.82 04s

Pisid44k 457 17 0.81 0.86

Plgid4s 13} 140 0.84 0.54

PIGtIn 158, 116 OTe OAS

P6167 152 116 0.98 (55

T16b 1.50 112 O88 56

Liodis children

Python spletes

Python umethtstinus

(er

Aspidives melonacephates

Chondropythan tiridiy

$9141.77 (182 +)

148-1,71 (1,60 + 02)

153-177 (1,66 oe 03)

4.74-1.88 (1.81 + 102)

1,671.76 (1-722 01)

(59-12-68 (1,62 + 01)

1,62-L.74 (167+ 01)

153-163 (158 + 91)

LA3-L48 (134 ye 097

OF8-305 CW Md)

OS1-1.10 (1.01 + 02)

LOI-ad (1.0 + 1)

VN41.25 11,204 th)

V.16-1.21 (V.18 e AS)

HAGEL LA (1 12 + 005)

TMB-11Z (109 01)

0:99-1.0% (Lod + 111)

D.75-0,99 (0.91 + 108)

O,77°LQ1 (OME + 2)

C7642 (Rd ON)

O75-40.96 (AS U2)

O.9S-1,08 (0,99 2 1)

O.3H-D.90. (0.89. 004)

0,760.83 (0.40 + 1)

M6947 (RA + 02)

1715-0,.82 (7B + 01)

O75-0,83 (0.99 + .01)

0.34-0.46 (40 > OL)

O.404).49 (0.45 + HN)

HAS (OSH A)

(540,59 (0.56 4. 01}

0.480.539 (O51 + 105)

O.41-4.45 (WAR + 005)

O.4241.47) (0.45 a= GON)

OATH. (0.42 11)

0.27-0.43 (0.37 01)

a gS

*Speciinen incomplete: yoty sbaot ene quarter of the precloacal vertebrae were studied.

TABLE 4

Length aid bropertions of dorsal vertebrae of specimens of three speeiva uf Varaous. The ulltinate presacral veeiebea uf euch specimen is exchuled The range Js follawed by

mean + standard ceror in purenihesis,

Pro (znm)

BwW/ Proto

CWlPCPa

Pelr/ ero

Yr BW

25-271 (25,7 + 14)

0.54-0:64 (0,58 + .005)

12.4-13,6 (13.14.07)

17.5-(9,.6 (18.5 + A)

147-15.8 (15.4 + .017)

0,48-0,60 (0.52 + .005)

047-059 (0.55 + 007)

0.43-0.54 (049 + sh)

0.35-0,63 (0.59 + 406)

052-056 (0.54 + .002)

0.48-0,.54 (1.52 = M4)

0.440.593 (0.49 =p N06)

0.88-1,01 (0,92 — O07)

.86-1,00° (0.90 4.007)

0.85-1.02 (0,97 + 00%)

O.82-0.95 (0.91 te DUT)

15i-f64 (1,58 + 008)

1.6641,78 (1.73 + .006)

1.70-1,68 (4,77 + 011)

17041,97 (4,65 + 013)

16,7418,.5 (179 +09)

13.0.15.4 (Lith te 1)

21,2-24.7 (2.9 17)

484059 (651 + 005)

OASOAT (SL + 007)

OAT. (OLS) OH)

0.56-0.59 (OSH + 002)

USIALS7 (1,55 E004)

At (0.49 + OO)

0.87092 (0,90 + .004)

0840.96 (091 + OH)

FER2-1AH) (U9 O11)

4.61-1.85 (1,76 + 012)

1,70-1.86 (1,78 + 011)

157-164 (1,73 + O15)

Numbers

Specimen of vertebrae

¥, gigantens 20

gould i:

Specimen t 20

Sneclmon 2 20

Specie S 2

VY, waerlies

Specinrert | (9

Speelmen 2 16

Specimen & y

¥ varius

{fornil) u

131-214 {163 + 0.67)

047-0455 (051 + 007)

U.48-0.57 (052 O07)

0.844), 98 (091 + O10)

1.65-1.37 (1.78 + 013)

—_——

SMALL FOSSIT. REPTILES FROM NARACOORLE 4u

physes or to the condyle} than in T. scincoides.

In Tiligua species and Evernia species the tip

of the neural spine may be thickened and some-

times marked with a shallow median grnove,

bucin 7. nigrolvrea (four specimens crammed)

the median grovve ts so deep that the spine

terninates in a double tip,

Three incomplete detitarics (P16124z,

P16126w, and PIG)28h) resemble 7. #fera-

lutea dentaries In shape, and four tmaxillac

(P16125s, P16128n, P16128w and P161572)

ore consistent with T. nierolutea (and atso with

T. scinvoides and 7. ovcipitaliv) in the slight

suborbital ridge. A parietal bone (P16127d) Is

probably also of this species although the sides

are slightly less constricted than in modern T.

aivrolutea.

cf, Spheaomorphus tympanum (Lonobery &

Andersson)

Material; Two fased sacral vertebrae,

In Sphenomorphus tympanum, the transverse

processes of the first sacral vertebrae (SI) are

strong and slant backwards only a few degrees.

The transverse processes of S2 are thinner and

are directed forwards to join and fuse with

those of Si at their lateral expunsions. The

sacral fossae between the transverse processes

are wide, §. ¢ympenut sacral yerlebrac differ

from those of Egernia stefofara in having tela-

tively wide fossae, and differ from EF. whirei

tenehrosa Where the transverse processes of S|

are angled backwards and the transverse pro-

cesses of S2 are perpendicular to the long axis

af the vertebra, ‘Tha fossil (P16146r) has a

total length (from prezygapophysis of Si to

posizygapophysis of $2) of 3,7 mm.

Egernia, c.f, E. whitei (I acepede)

Material: V ertebrace—dorsal (2, lengths 3.4, 2.7

mm), caudal (1, length 2.4 mm); manillac (5

Icft, 5 right); dentarics (8 lett, 6 right); fron-

tals (1).

The Meckelian groove in the lower jaw is

closed anlerior to the splenial in Egernia but it

is open forward to the symphysis in Spheno-

tntorpius. Vhe detitary of EB. whitei is deeper

than the slender dentary of S$. gvinpanien, and

the notch in the posterior lateral surface of

the dentary is higher (i.e. nearer the tooth row)

than in S. lyenpanunt, The fossils wre consis-

tent in shape and size with E. whirei,

The fused frontal bones of E, whitei differ

from those of §. ryepanuen in their gradual

taper, both anteriorly and postenurty.

Family AGAMIDAE

Agamid vertebrae ale characterized by therr

trangular Ventral outline and slrong subventral

ridges.

Amphibolsrus c.f, A. barbatus

Material: Maxillac (1); dentaries (7).

Agamids ate (he only Australiqn reptiles with

acrodont tooth implantation. The largest fossil,

a right dentary (P16132b) with length af tooth

row 14.5 mm, closely resembles 4. barhatiy.

The other specimens, two of them fragments,

may he of a smaller specics,

Faunal change

The two soptiles most common in the Victoria

Cave deposit, viz. Pseudonaja c.f, Po auehalis

and 7. ragosas were represented at all depths

in similar abundance. The less common species,

except for #4’. neracoofiensis were also found

al various depths from the surface to the

present maximum depth of excavation, Sevei

vertebrae und the tooth fragment of MW, nare-

coortensis Were near the surface and all within

2 metres of each other. Hence the reptile fauna

does not change remarkably with depth in the

deposit.

Discussion

The small marsupial remains, together with

ubundant rodent vemains, were probubly

brought into Victoria Cave by owls (Smith

1971, 1972), and the small lizards may also

have been the prey of owls, Among the larger

spovies, Trachydasmirus rugosus is a clumsy,

shortlegyed, heavy-bodied livard which might

easily fall into sinkholes or caves and would

have liltle chance of escaping, This species has

been recorded from several cave deposits (c.g.

Cook 1963, Finlayson 1933, Longman 1945),

The snakes may have actually inhabited the

cave, as live brown snakes (Psendonaja sp.)

ave found in the limestone caves in south

eastern South Australia (Wells, pers, comm,)

and P. auchalis has been classified as an occa-

sional troglonene (Richards 1971),

Tn any measurements af the bones of rep

tiles, intraspecific Variances are large because

reptile growth is asymptotic. When vertebrae

ate the bones measured, changes along the

column further increase the variation. Jn the

identification of isolated vertebrae of some

groups (eg. the snake family Crotalidac),

these inhetent large variances can he offset by

considering several dimensions simultaneously

and in comparison with their previously deter

mined inter-relutionships along the entire

columo of reference skeletons (Brattytrom

40 MEREDITH J. SMITH

1964). Nevertheless, when the reptile remains

ure abundant or include « qualitatively diag-

nostic bone (ee the first sacral vertchra of

Vurunus), the specics can be diagnosed with

cunfidence,

Of the 3 species contidently determined, 4

are still Jound in south-easiern Australis, antl

all but Faraaus varius have been found near

Naracoorte. All 6 additional species tentatively

identified have been found near Naracoorte.

The large boid is the only Pleistocene species

absent now, Hence, among the reptiles, the

small species have survived from Pleistocene to

presem without detectable change of the

vharacters available in fossil muterial, whereas

the large species has become extinct, Similarly

with the marsupials; while many large species

have become extinct feg. several Sthemrus

species, Thylacelea «ff. 7. carnifex (Wells,

pers. comm,), Palorchestes sp., (Pledge, pers.

comm.,)], the small species, ¢.g. Berrengia spp.,

Perameles spp, Antechinus spp. and Perauras

breviceps are indistinguishable from modern

species, many of which suill survive near Nara-

eoorte (Smith 1971, 1972), Among the birds.

the only specics now exunct. Progra nara-

coortensix, Was a large bird, while all of the

small spectes ire extant. The factors that

caused the extinction Of so many large verte-

brate specics have had little perceivable effect

on the small vertebrates.

The presehee of Paranus varies together

with V, goulii? in Chis Pleistocene deposit does

Hol support the suggestion (King & King 1975)

(hut the iedicus karyotype (represented by 1.

varius) tavaded south-eastern Australia after

lhe separation of Kangaroo Island from the

mainland, 3,000-10,000 years ago.

Most of the extant species of the Vicloria

Cave seplile fauiny are wide-ranging with broad

habitat tolerances, Varanns gouldii occurs in

most parts of mamnland Australia but is most

commen in sandy areas, where it lives in sand

burrows. (Worrell 1963). The tree-climbing

species WM. varius, otcurs throughout exstern

Australia inside the 20” (508 mm) isohyct

(Rawlinson 1969), Trachydosaurus rugusus. is

found in inland areas of all muinland stutes,

while Pserdechix porphyriacuy lives in coastal

to mountainous forests and swanips of eastern

Australia, but does: not extend into dry inlani

areas (Worrell 1963). The ranges of Pseuda-

taja nuchalis and ihe marphologically similar

P. rextilis together include most of mainland

Australia (Worrell 1963), and P. fextilis o¢ours

also in New Guinea (McDowell 1967). None

of these species extend inte the cool temperate

zone of the Bassian zoogcographical subregion

(Rawlinson 1974). Conversely, Tiliqua nigro~

lures is contined to the cool temperate zone, its

Tange extending from the extreme south-casi

of South Australia and southern, Victoria to the

islands of Bass Strait and Tasmania. Nura-

coorle is close to the north-western limil of its

Tange (Rawlinson 1974). Evernia = witiret.

Sphenomorphus tympanum and the genus

Notechis occur in all zones of the Bassisn but

not in other subregions (Rawlmson 1974)

Hence little palaeo-ecological infonmution can

be gleaned from them, The presence of a lirve

Proportion of the Pleistocene reptile fauna i

the area at present does suggest that climatic

changes during the last 30,000 years have been

slight in south-easicrn South Australia.

Acknowledgments

The field work im Victoria Cuve has heen

planned and supervised by Dr R, T, Wells, and

to him and his voluntary helpers (many of

them CEGSA members) I am grateful. Trans-

port costs for field workers were defrayed by

a gram from the South Australian Government

Tourist Bureau. Specimens of snakes and

lizards have been given or loaned to me by

many pedple, among Whom I thank especially

Mr F. W. Aslin, Mr J. deBavay, Mr W. J.

Parmenter, Mr R, Shine. Mr M. J, Tyler, Dr

R. Henzell. Dr BD. Harton, Dr T, F, Houston,

and Dr G. F. van Tets. Mr P. G. Kernpster

prepared the photographs (Fig. 2), Dr R, T

Wells and Mr M. J. Tyler kindly criticized the

rhanuscript.

References

ANbrews, C. W. (1906).—"A descriptive

catalogue of the Tertiary Vertebrata of the

Fayum. Exypt.” (British Musgenm: London.)

Ancein, M. (1974).—Excavations in the Orches-

tra Shell Cave, Wanneroo, Western Australie.

Fl, Fossil vertebrate tenmains, Archeol, Phys.

Anihrapal, Oceana 92), 156-162.

ASTEFENBERG, W. (1963 —The lossil snakes of

Florida. Tulane Sid. Zool, 1, 131-216

Bedrtstrom, B. H, (1964),—Eyalution of the pit

vipers, Frans, Sun Diezo Soc. Nat. Hist.

13( 11). 185-268-

Coon, D, T- (1963)—The fossil vertebrate fauus

of Strone’s Cave, Koranup. Western Australin.

W, Aust, Nat. %, (53-162,

Dortcr, C_ E,. & Mrrarirens, D, (1971).—A sal-

vate excavation in Devil's Lair, Western Ats-

tralia, J, R. Soc, W, Ast, 54, 103-114.

SMALL FOSSIL REPTILES FROM NARACOORTE 51

Epwusp, A. G. (1969)—Dentition. In C. Gans

(Ed.), “Biology of the Reptilia”, Val- [, pp.

117-200. (Academic Press: London, }

iain, A. P. (1964).—The Anstralian Aborigines.

4th edn. (Angus and Robertson: Sydney.)

Frinavary, G. J. (t918)—Contributions io a

monograph on fossil Yaranidac and on

Mesgwlanidae. Ann, Aist, Nat. Mus. Nail

Hung, 16, 341-467.

Pesrrvary, G, J. (1935) —Forther contributions

to a monograph of the Megalanidue and

fossil Vurunidae. with notes on recent

Varaniany, Ann, Aisi. Nat, Mus. Nat! Hung.

29, 1-130.

Fincayson, H. H. (1933). dn Tindale, N, B.

(1933).—Tantinoola Cuves, south east of

South Australia: geological and physiographi-

cal notes. Trans, R, Soc. 8S. Aust, 57, 130-142.

Gre, B,D, & Banns, M, R. (1956),—Cainozoic

history of Mowbray Swamp and other areas

af north western Tasmania, Rec, Queen Vic-

fori Mus. Launceston, N.S. No. 6, 1-42.

Hecut, M. K, (19753)—The morphology and

relationships of ihe largest known terrestrial

lizard. Megalania prisca Owen, fom the

Pleistocene of Australia. Proc. R, Soe. Viet,

87(2). 259-250,

Horrstetrer, R. (19610),—Nouvegux restes d'un

serpent boide (Madstoia madagasceriensis

nov. sp.) dans le Cretace supericur de Mada-

gascar. Bull, Mus. natl Hist. nat., Pariy (2),

33, 152-160,

Horrstetter, R. (1961b).—Novvelles recoltes de

serpents. fossiles. dans I‘Eocene Superieur de

Deseri Libygue. Bull. Mas. natl Hist, net.,

Paris (2), 33, 326-331.

Tlorrsvettre, R., & Gase, -P (1969). —Vertebruc

and ribs of modern teptiles, 4a C, Gans (Ed).

Biology of the Reptilia Vol. I, pp. 2016310.

(Academie Press: Loyidor.)

Horrstetiin, R.. & Gavara, ¥. | 1964) —Obser-

vations sur l’osteologie et ta classification des

Achrochordidae (Serpentes). Ball. Mas. nail

Hist. nat,, Paris (2) 36, 677-696.

HeEnsitkoviTz, P. (1972).—The recent mannmnals

of the Neotropical Region: A zoogcographic

und ecologival review. Jn A. Keust, F.C. Erk,

& B. Glass (Eds), “Evolution, Mammals and

Southern Continents.” (State University of

New York Press: Albany.)

King, M,, & Kina, D, (1979) —Chromosomal

evolution in the lizard venus Maranns (Rep-

lilia), Aust. J. Biol. Ses 28(71), 89-108,

Loncman, H. (1925)—Ophidian vertebrae

tram cave deposils al Marmor Quarry, Meni,

Old Afus, 8, 114-112,

Lonoman, H, A, (1945)—Foasil vertebrates from

Gore Quarries. Mem. Qld Mus. 12, 164

Lunpriaus. E. L. (1963).—Vertebrate remuint

from the Nullarbor Caves, Western Austrulin.

J. R. Sac. W. Aust. 46, 75-80.

Lyprekéa, R. (1888),—“Catalogue of Fossil Rep-

tiles and Amphibians" Part L (British

Museum: London.)

McDowe1s, SB. L1967}—Aspidomorphus, a

genus of New Guinen snakes of the family

Elapicdae, with notes on related genera. J.

Zoel., Lond. 151, 497-543.

MesRILEES, D. (1968).—Man the- destroyer: Tate

Quaternary changes in the Australian mar

sugial fauna. J. R, Soc. AW’, Aust, 51, 1-24.

Mertens, R. (1942),—Die Familie der Warane

(Varanidae). Zweiter Teil: Der Schadel, Abh,

Senckenb. Nutarf, Ges, 465, 117-234,

MrreHett, F. J, (1950)—The scincid genera

Egernia aod Tiligua (Lacertilia}. Ree. $. Ass.

Mas. 9, 275-308.

Rawranson, P. A. {J969).—The reptiles of Exsr

Gippsland, Proce. R. Svc. Piet. 82, 113-128,

RAWLINSON, P. A. (1974).—Biogcography and

eculogy of the reptiles of Tasmania and the

Bass Strait area. Jn Williums, W. D, (Ed.),

“Biogeography and Ecology in Tasmania’, pp.

291-338. (Junk: The Hague.)

RacHagus, A. M, (1971),—An ecological study of

the cavernicolous fauna of the Nullarbor

Plain, South Australia, 7, Zeal. Lod, 164,

1-60.

Romer, A, S, (1956),—“Osteclogy of (he rep-

tiles” (University of Chicago Press:

Chicage.)

Simpson, G. G. (1933) —A new ossil snuke from

the Nofastvlops beds of Patagonia. Bull. Arm.

Mus. Nat Hiss. 67, 1-22.

Surea, M. J. (197)).—Small fossil vertebrates

from Victoria Cave, Naracoorte, South Aus:

tralia. [. Potoroinae (Macropodidae), Petau-

ridac and Burramyidae (Marsupialia)., Trav.

R. Sov. 5. Aust. 95(4), 185-198.

Suen, M. J, (1972),—Smull fossil vertebrates

from Victoria Cave, Naracoorte, South Aus-

tralia. IT. Peramelidac, Thylacinidse ancl

Dasyuridae (Marsupiulia). Trans. R, Sar. 8.

Aust, 96(3), 125«137,.

Suity, M. J. (1975).— The vertebrae of Four Aus-

tralian elapid snakes (Squamats; Elapidae).

Trans. R. Soc. 8. Aust, 99(2), 71-84,

Trorwe, A. (1971)--The Fauna. In R, Vo S

Wright (Ed.4, “Archseolory of the Gallus

site, Koonakky Cuve™ Australian Inst.

Aboriginal Stinlics, Na. 26, 45-47,

VAN TrTs. G. FP, & Sark, M. I} (1974).—Small

Fossil vertebrates Froni Victoria Cave, Nata-

cuurte. South Australia Til, Birds (Aves).

Tras, R, Soe S$. Aust. 9B(4), 228-227.

Worrett, £. (1963}—"Reptiles af Ausititra’,

(Anges and Robertson: Sydney.)

VOL. 100, PART 2 31 MAY, 1976

TRANSACTIONS OF THE

ROYAL SOCIETY

OF SOUTH AUSTRALIA

INCORPORATED

CONTENTS

Archer, M. Miocene Marsupicarnivores (Marsupialia) from Central South

Australia, Ankotarinja tirarensis gen. et sp. nov., Keeuna wood-

burnei gen. et sp. nov., and their Significance in Terms of Early

Marsupial Radiations - - - - > - - - 53

Reedman, D. J., and Womersley, H. B. S. Southern Australian species of

Champia and Chylocladia (Rhodymeniales: Rhodophyta) - 75

Watts, C.H.S. Leggadina lakedownensis, a New Species ef iad Rodent

from North Queensland - - - - - 105

Baverstock, P. R., Hogarth, J. T., Cole, S., and Covacevich, J. Biochemical and

Karyotypic Evidence for the Specific Status of the Rodent

Leggadina lakedownensis Watts - - - - - - 109

PUBLISHED AND SOLD AT THE SOCIETY’S ROOMS

STATE LIBRARY BUILDING

NORTH TERRACE, ADELAIDE, S.A. 5900

MIOCENE MARSUPICARNIVORES (MARSUPIALIA) FROM CENTRAL

SOUTH AUSTRALIA, ANKOTARINJA TIRARENSIS GEN, ET SP. NOV.,

KEEUNA WOODBURNEI GEN. ET SP. NOV., AND THEIR SIGNIFICANCE

IN TERMS OF EARLY MARSUPIAL RADIATIONS

BY M. ARCHER*

Summary

ARCHER, M., (1976) .-Miocene marsupicarnivores (Marsupialia) from central South Australia,

Akotarinja tirarensis gen. et sp. nov., Keeuna woodburnei gen et sp. nov., and their significance in

terms of early marsupial radiations. Trans. R. Soc. S. Aust., 100(2), 53-73, 31 May, 1976.

Two of Australia's oldest known marsupicarnivores, from the Etadunna Formation of the

Lake Eyre Basin, are described. Ankotarinja tirarensis is a tiny marsupicarnivore which may be

related to didelphids as well as dasyurids. Although it is much too late in time to be the actual

ancestral dasyurid, it is regarded as a structural ancestor. It is also structurally ancestral to Keeuna

woodburnei.

Keeuna woodburnei is a small marsupicarnivore which is more similar to dasyurids than is A.

tirarensis. It resembles species of Phascolosorex Matschie, 1916, Neophascogale Stein, 1933,

Murexia Tate & Archbold, 1937, and some Antechinus Macleay, 1841. More distant relationship to

didelphids is suggested.

Resemblance of both of these fossil marsupicarnivores to modern New Guinean highland rainforest

dasyurids rather than to more arid-adapted Australian dasyurids, is regarded as evidence suggesting

that central Australia was less arid during Etadunna time than it is now.

MIOCENE MARSUPICARNIVORES (MARSUPIALIA) FROM CENTRAL SOUTH

AUSTRALIA, ANKOTARINIA TIRARENSIS GEN, ET SP. NOV., KEEUNA

WOODBURNEI GEN. ET SP. NOV., AND THEIR SIGNIFICANCE IN

TERMS OF EARLY MARSUPIAL. RADIATIONS

by M. ARCHER”

Summary

ARCHER, M., (1976),.—Miocene marsupicarnivores (Marsupialia) from: central South Australia,

Akotarinja trarensis gon, ct sp, nov., Keeuna woodburnel gen et sp. nov,, and thelr

sigttificance tf terms of early marsupial radiations. Trans. R. Sec. S. Anst.,100¢€2), §3-75,

31 May, 1976,

Two of Australia's oldest known marsupicarnivores, from the Etadunna Formation of the

Lake Fyre Basin, are described, Ankotarinja tirarensis is a tiny marsupicatnivore which mity

be related to didelphids as well as dasyurids. Although it is much too Jate in time to be the

actual ancestral dasyurid, it is regarded as a structural ancestor, It is also structurally ancestral

to Kéeeuna woodburnei.

Keeuna woodburnei is a small marsupicarnivore which is more similar to dasyurids than

is A. tirarensiv. Tt resembles species of Phascolosorex Matschie, 1916, Neophascogale Stein.

1933, Murexia ‘Tate & Archbold, 1937, and some Antechinus Macleay, 1841, Mote distant

relationship: io didelphids is suggested.

Resemblance of both of these fossil marsupicarnivores to modern New Guinean highland

raiiforest dasyurids rather than to more arid-adapicd Australian dasyurids, is regarded as

evidence suggesting that central Anstralia was less arid during Etadunna time than it is now.

Introduction

In 1971 W, A. Clemens, M, 0, Wood-

burne, C. Campbell and the author recovered

fossil] mammal remains from a site known as

Tedford Locality, on the west side of Lake

Palankarinna, Etadunna Station, South Austra-

lia. These fossils come from the Etadunna For-

mation which is now regarded (W. K. Harris,

pers. comm.) ys being approximately middle

Miocene in age. In 1972 M. O. Woodburne, P.

Lawson, W, Head, B. Archer and the author

extensively quarried and screen-washed Ted-

ford Locality. From the concentrate, two mar-

supicarniyores, as well as other mammal re-

Mains, Were Tecovered.

Stirton, Tedford, & Miller (1961) briefiy

describe a third marsupicamivore Irom the

Etadunna Formation.

Terminology of individual tceth is that used

by Thomas (1887) and Archer (1974). Ter-

minology of tooth crowns is shown in Figure |

and follows that used by Archer (1975a),

Comparisons With other marsupicarnivores are

based in large part on Archer (1976b}.

Specimens with prefix P are in the fossil?

collection of the South Australian Museum; F

in the fossil collection of the Queensland

Museum; UCR in the University of California

at Riverside; AMNH in the Archbold Callec-

tions of the American Museum of Natural His

tory; J and JM in the modern collections of the

Queensland Museum; and WAM in the modern

collections of the Western Australian Museum,

Species names of modern Australian niar-

supials are those employed by Ride (1970),

Laurie & Hill (1954) or Archer (1975b).

Other modern marsupial pames arc those

employed by Collins (1974), Names of Creta-

ceous didelphids are mainly those used by

Clemens (1966). Fassil marsupial names are

those. employed by the most revent reviewer of

those particular groups.

Taxonomy

Genus ANKOTARINIA nov.

Type species: Ankotarinja tirarensis sp. nov,

(by designation and monotypy).

Generic diagnosis; Differs from other Austra-

lian and New Guinean dasyurids in haying, as

* Queensland Museum, Gregory Terrace, Brisbune, Old 4006,

54 M. ARCHER

~ pe

Fig. }. Terminology of molar cusps and crests

(based on Ankotarinja tirarensis). 1A,

upper molar. 18, lower molar, a,c), an-

terior cingulum; c.6,, gristid. obliqua; end,

entoconid; Aycd, hypocristid; hyd, hypo-

conid; Ayld, kypoconulid; hyld n., hypo-

conulid notch; mel., metaconule; melr.,

metaconular ridge; me., metacane: nec,

meticrista; mecd, metacristid; med, mets-

conid; pa., paracone; pacd, paracristid;

pad, paraconid; panie. cr., para-melacone

crest; pel., protoconule; pprer,, postproto-

crista; prer,, protocrista; prd, pratotonid,;

pred, precingulid (or anterior cingulum);

psed, postcingulid {or posterior cingu-

lum); pasted, parastylid; r., ridge mesial to

st©; stA-E, stylar cusps A-E.

a combination of characters, relatively wnre-

duced tafanid on M4 with well-formed hypo-

conid, hypoconulid and entoconid and rela-

tively unrecduced P4.

Origin of generic name: An allusion to Anko-

tarinje, a dreamtime ancestor (Robinson 1966,

Pp. 26) who, having remained buricd a long

tinmte as bones in the earth, resurrected himself

in a small watercourse. Asikorarinja is here

given masculine gender.

Ankotarinja tirarensis sp. nov.

FIGS 1-5

Holotype, P18190, right dentary fragment with

MS.

Type locality: Tedford Locality, Etadunna For-

mation, Lake Palankarinnta, Etadutma Station,

S.A. (28°47'S, 138°25'B).

Diagnosis; That of genus. Features likely to be

of specific value include very small size, rela-

lionship of hypocristid to entoconid, size and

width of anterior and posterior cingula, relative

size of paraconid on M4, and relative size of

stylar cusps,

Origin of specific name; Specific name refers to

the Tirari Desert, the portion of the Simpson

Desert-containing Lake Palankarinna.

Referred specimens: UCR, 15340, dentary

fragment with LM4; UCR, 15341, dentary

fragment with LMs-4; UCR, 15342, dentary

fragment with RM4; P7331, dentary fragment

with LM+-4; UCR, 15343, maxillary fragment

with alveoli for LM3-4; UCR, 15308, LM**;

F7332, LM?3.

Description

Maxillary fragment (UCR, 15343) referred

to this species on basis of size, has alveoli for

M8-4 and posterior root of M2. M4 appears ta

have been as wide as M34, presumably with

relatively little reduction of protocone. M4

length less than that of M#, metacone roat

heing veduced and displaced antero-lingually

relative to metacone root of M%, Zygomatic

root of maxilla yrises buceal to region between

M3 and M4. Numerous smal) interdental

fenestrae in palate hetween M3 and M3, and

between M3 and M4,

Upper teeth represented by two isolated

molars, probably LM% and LM+4. Although

possible that these teeth actually represent Mi},

und M3, improbable for following reasons:

Stylar cusps (UCR, 15308) much more

reduced than those cusps on M®& in all other

dasyurids but not so strikingly reduced when

compared with M4 in some dasyurids such as

Keeuna, described helow, or some species of

Planigale Troughton, 1928, Also, notch tm

antero-huccal cingulum of F7332 for reception

of metaslylar corner vf preceeding tooth sug-

gests F7332 is posterior molar. However, it is

also true that in some dasyurids and didelphids

with large P+, such a notch sometimes exists

MIOCENE. MARSUPICARNIVORES a8

Fig. 2. Specimens of Ankerarinja tirarensis and their measurements (mm). 4, F7331, LMé-4. & UCR.

15340, LM4. C, UCR, 15341, LMs-, BD, Holotype, P18190, RM{4. E, F7332, LMS. F, UCR,

15308, IMé G, UCR, 15343, maxillary fragment with alveoli for | M24.

in antero-buccal cingulum of M1. Probable

that F7332 and UCR. 15308 represent M2 and

M4 respectively rather than M4 and Ms.

F7332 has at least three stylar cusps. StB

connects to parastylar corner of tooth which

may be distinguishable as discreet stA, Para-

stylar blade very short. Ectoloph crest descends

gently from posterior flank stB, then rises

gently, to form long low ridge-like stC, Pos-

tenor paint of this cusp marked by beginning

of rise in ectoloph which forms tall stD. 5iD

has minor ectoloph crest on posterior flank

which descends towards metastylar corner of

toath. Very minute rise in crest on posterior

flank of stD may represent stE, Ectoloph pos-

terior to this point very low, extending to meta-

stylar corner of tooth. Paracone taller than

stylar cusps but subequal in crown height to,

or shorter than, protocone. Mefaconc tallest

cusp. Prominent protoconule and metaconule.

Ectoloph continuous on buccal edge of crown.

Buccal concavity in crown outline slight. Para-

¢rista just longer than half length metacrista.

Paracrista extends from paracone to anterior

half of stB, Although slightly worn, paracrista

appears to curve at buccal end to contact stB.

Paracrista apparently transverse to imaginary

long axis of toothrow. Para-nictucrista. con-

tmuous, Slight protoconule ridge may be

present linking base of paracone to preproto-

crista, Clear metaconular ridge present linking

base of metacone to postprotocrista. Meta-

conular ridge extends short way up base of

Metacone causing bulge in base of that cusp,

Metaconular ridge bounds marked declivity

between posterior portion of steeply inclined

postprotocrista and posterior base of metacone.

Anterior cingulum complete, linking preproto-

crista and antero-buccal cingulum to parastylar

comer of tooth. No posterior cingulum present.

Pre- and postprotocrista form large but acuie

angle,

UCR, 15308 has at least four stylar cusps.

Possible stA as in F7332. Posterior crest from

stB descends steeply to base of stC. Between

stC and D, and connected by crests, another

smaller stylar cusp of uncertain homology. StI

small and connected to metastylar corner of

Sf M, ARCHER

tooth by low ecteloph crest, Paracone and

protecone subequal in crown height. Prote-

comple absent and protacenule swelling only

Just present. Metaconule large. Buccal con-

cavity in crown outline deeper than in F7332.

Paracrista almost three-quarters length meta-

crista, Puracrista wom but appears to intersect

ectoloph ow anterior flank of siB. No clear

protoconule ridge present, Metaconular ridge

sovall bur present, Metaconular ridge docs apt

clearly extend up base of metacone, Otherwise

morphology of UCR, 15308 as.in F7332.

Meristic gradients from M5 to M4 may be

summarized as follows, Tooth length decreases

Width increases, Ectoloph becomes more

evenly and decply concave. Stylar cusps, par-

licularly OD, become smaller. Paracristy atid

metacrista Increase in lenvth. Para-metacrista

hecomes more synitietrical. Prococemule de-

creases in size. Angle formed by pre- and post-

protocrista hecormes slightly more acute.

Antero-buccal cingwlim increases stghtly in

tength,

Dentary fragments Indicate premolar size.

UCR, 15340 hus alveoli for C4-P4- Premolar

alveoli subequal in size, indicating lite or no

reduction in tooth size fram Pi ta P4- Py, pre-

sumably had posterior cingular cusp which

corresponds with hypocornulid notch of My_ Py

anterior alveolus slightly crowded out of align-

ment but (as evidenced in modern species of

Plonigale, Accher |976a), does not necessarily

mean P44 crown gut of aligninent. Judging

from proximity of premolar and canine alveoli,

premolars and canine presumably contacted

one another antero-posteriosty. C4 alveolus

suggests Cy width exceeded that of any pre-

molar, but because of relatively unreduced Py,

C{ probably not greatly enlarged and com-

parable with canine of Nirgauw/ Archer, 1975b.

Mz? talonicd wader than inigonid but trigonid

not as compressed laterally as in most modern

dasyurids, Well-ieveloped anteniar cingulum,

which terminates Lingually for hypoconulid

notch, Parastyiid comer of tooth most anterior

portion of crown. Posterior cingulum com-

parable in Jength to anterior cingulum and ter-

minates buccal to posteriorly projecting hypo-

conulid, Basal cingulum absent beneath

postero-bireceal corner of protoconid and hypo-

conid. Roughened enamel suggests cingulum

present hetween base of protoconid and hypn-

conid, No lingual cmgulum, Paraconid low,

approximately same height as hypoconulid.

Protoconid tallest cusp of irigonid, Metaconid

just shorter thin proteconid, Hypovonid just

taller than entoconid which is taller thai pare-

conid, Paracristid complete between protaconid

and paraconid but almost vertical from proto-

coaid to shallow paracristid fissure and hor)

zontal between paracristid fissure and para-

conid. Metacristid steeply inclined on both

sides of metacristid fissure. Metacristid und

hypoeristid approximately transverse to long

axis of dentary. Cristid obliqua (damaged)

extends from hypoconid to trigonid intersecting

latter at point below protoconid tip, well buccal

to metacristid fissure. Hypocristid extends From

hypoconid to hypoconulid, without approach-

ing entoconid. Entoconid and hypoconulid net

connected by crest. Entoconid and metaconid

connected by high crest.

Ms talonid wider than trigonid. Antertor

and postenor cingulum as in Mj. Buccal cing

lum helween protovonid and hypaconid Jess

developed (absent in P18190). No lingual

cingulum, Paraconid smallest trigonid cusp but

subequal in height to hypocenid and entoconid.

Melaconid much taller than hypoconid and

just shorter than protoconid, Hypoconid and

entoconid subequal in height. Entoconid not

connected ta hypoconulid hy crest, but con-

nected to metaconid as in M4 except that crest

interrupted by shallow transverse groove, Para-

cristid from paracristid fissure ta paraconidl, in-

clined, nor horizontal, Crista obhqua extends

to base of protoconid as in My; but anterio¢

end appears tu be distinct contribution from

trigonid with slight fissure where talonid and

trigonid parts meet. Trigonid portion thicker

and more bulbous. Otherwise morphology Ms,

asin Mg.

Mg talonid just wider than trigonid. Anterior

and posterior cingulum as in Ms. Buccal cingu-

lum confined to arca betwecn base of proto-

conid and hypoconid, as thickened bulze of

enamel, clearly less well-developed than

anterioy and posterior cingula, No lingual

cingulum. Paraconid smallest trigonid cusp but

taller than any tabonid cusp. Entoconid not

connected ta hypoconulid hy crest, but con-

nected to metaconid, as in MG. Unlike eristid

obhiqua in Ma, this structure in My appears to

lack transverse fissure separating ¢rest inte

hypoconid and trigonid portions: This dif

ference between Mi and Ms, notable in

P18190, Cristid obliqua, also intersects trigonid

in slightly more lingual position than m Ma,

Otherwise morphology M%, as in M4.

My trigonid wider than talonid, but talonid

wider than that stricture in most modern

dasyurids. Anterior cingulum as in Ms, Pos

MIUCENE MARSUPICARNIVORES AF

terior cingulum absent. Buccal cingulum can-

fined to arta between protoconid and hypo-

conid, Paraconid just shorter than metaconid.

Etitoconid and hypoconid reduced relative to

My, but larger relative to most modern

dasyurids. Entoconid connected to base of

metaconid via low crest, Entoconid also com-

nected to Aypoconulid by low crest, Hypo-

crshd convex anteriorly. Hypoconulid sub-

equal in height to entoconid. Cristid obliqua in-

tersects trigonid base immediately buccal tu

point below metacristid fissure, this being

markedly more lingual than similar intersection

of cristid obliqua in My. Otherwise morphology

of M4 2 in Mg.

Meristic changes along touth row as follows.

Payuconid increases in height from M{ to Mj.

Melaconid height Mj4-4 subequal but meta-

cond length at byse of cusp decreases

tnackedly from M4 to M4, result of reduction

In size @f minor crest on posterior slope of

metaconid which links with enteconid, Ento-

conid M;- subequal and larger than ento-

conic of Mj, Hypoconulid M4-4 subequal in

height aud shorter than that cusp in Mi‘. Proto-

conid M4 shorter than protovonid My which

is subequal 10 that cusp in M4 which is larger

ihan that cusp it M4. Hypoconid decreases in

height from M4 ta Mg, Talonid M4-4 wider

than tigonid. Talonid M4 narrower than

trigonid. Paracristid M4-4 subequal and sub-

equal lo (P18190) or just shorter than (UCR,

15341) that crest in Mg. Paracristid M4

shorter than that of M4. Metacristid increases

in length from M4, to Mé. Metacristid My

shorter than metacristid M4. Cristid obliqua in-

tersects trigonid in progressively more lingual

Position from Mi to M4. Hypocristid My -{

subequal in length and longer than hypocristid

My. Anterior cingulum decreases in length

slightly from M4-M4. Posterior cingulum

M4 -:, subequal in length (absent in Mi)

Discussion und comparison

Anketarinja t a Metatherian because it has

four molars, a larec stylar shelf, and an

approximaled entocenid and hypoconulid. It

is also a marsupicarnivore because it has tribo

sphenic molars lacking hypocencs,

Denta| characietistics of known dasyurids,

peramelids, thylacinids, didelphids, and related

marsupicarnivores have been reviewed

(Archer 1976b) and, to avoid repetition; it is

sufficient to point out here that Avkerarinja

can only be regarded as either a dasyurid or

didelphid, Because morphological yariation of

teeth exhibited by didelphids and dasyurids

OVerlaps, only incisor number and possibly

dP4 cusp number permit diagnosis at the

family Jevel. Al) dasyurids have three lower

incisors and four upper jacisors on each aide,

whereas almost all didelphids have four lower

and five upper incisors (exceptions may include

Derorhynchas singularis Paula Couto, 1952,

species of Eodefphis Matthew, 1916). The pre-

maxilla and amlerior region of the dentary of

Ankotarinja Hrurensis are unknown, Therefore,

this marsupicarniyore cannot at present be

referred conclusively to either the Dasyuridae

or the Didelphidae,

Modern and fossil didelphid subfamilies

exhibit greater morphological variation than

dasyurids. Most are so distinct that their mere

subfamilial status has been questioned, and the

reasons they are doubtfully referred to the

Didclphidae are also the reasons they cannot

be related to Ankotarinija tirarensis. Only didel-

phines warrant closer comparison.

Some North and South Anictican didel-

phines are adequately illustrated (such as the

Paleocene forms described by Paula Couto

1952, 1962, 1970} but most are not. Archer

(19766) summarizes the must striking charac-

teristics of these forms, Only species of Coona

Simpson, 1938, Marmoyepsis Paula Couto,

1962, Mirandetheriunr Paula Couto, 1970,

Monadelphopsis Paula Couto, 1952, Derur-

hyachus Paula Cotte, 1952, [schyredidetphis

Paula Couto, 1952, Didelpieopris Paula Coute,

1952, Minwseulodelphis Paula Couto, 1962 and

an M+ listed by Paula Couto (1962) as incer-

fae sediv are similar to A, tirarensis, Among

Cretaceous didelphines, species of Alphadan

Simpson, 1927, and Pediowys Marsh, 1889,

warrant comparison. Modern didelphines used

here for companson include two species of

Marmosa Gray, i821, M. sp. and M-~ siitis

Bangs, 1898, Monodelphis dimidima (Wagner,

1847), Metachirus nudicaudanus ( Geoffroy,

1803), Philander opassun: Gray, 1843, and

Didelphis marsupialis Linnaeus, 1760, Of

these, species of Marmosa, Monedelpits Bur-

nett, 1830 and Didelphis Linmacus, 1758

warrant closer compucison with 4 nkofariila.

Characters of Ankotarinja which are wousual

among dasyurids and inVite broader compari

son within the Marsupicarnivera (the didel-

phine forms noted sbove) are as follaws: 1,

large M4 talonid; 2, relatively uncompressed

My trigonid: 3, huccal position of anterior end

of cristid obliqua; 4, transverse orientation of

metacristid: 5, large P4; 6, large M+; 7, stylar

jaa

Ss)

MIOCENE MARSUPICARNIVORES

psc

MIOCENE MARSUPICARNIVORES aL

TABLE =

= 22

z= =- 4

eisigde2Fs : Ei.

+ BEEPS EBE = = oc 5 = & =

= eSSesasgosesttEtses BS so = 3

a SStetPyVSEFrFsPsEerzsrTtSTrErzrsrzers te

23aé SEEPRBERQGFTSEBEPBRERESZ RE ESE Z

. SEESSSSSEFESSZSEESTEREER ERE =

Unusual characteristics $4 & Se iszeii =a TESPESS €£ 28 3

of Ankotarinia NHS SPTQRARSFS oS tre FBR SFeRrRzAA

1 Mj talonid Jarge + + F— tt 7 Lu, a4 tre gt ats b/ + ——— — 9

2% My trigonid wide ; Pa pt he F Bab ff BE feof fee =

3. Coo, buecal position +4 bope tee PHHeE F444 2 4 -——— H +

4. Transverse metacristid + }| | -—-—- -—-— Se

S. Large P4 + ele Pepe ole — -= -apepeaee tony —- 7

® Large Mg + YP 7 PP Fo PPE HHA HF HERK HE H HH HH |

7. stD small attd posterior Se re i a ey ec a ces eee ee +

6, Metaconule prominent FH FY He Veer MPR tem H tt ie eH Le 6b

cusp arrangement with reduced and posteriorly

positioned stD; and 8, prominent metaconule,

These characters are compared in Table 1 for

non-dasyurid marsupicarnivores noted above,

as well as dasyurids which provide the closest

approximations to Ankotarintja. In this Table,

characters 1-8 are the same as noted above.

A "+" records a condition similar to or which

approaches the condition found in Ankotarinja.

A “?" indicates either that the requisite part of

the dentition is unknown or not adequately pre-

served, A “—” indicates ‘a character condition

unlike that in Ankertarinia. In some genera

noted, some species may be similar to, while

others differ fram, the condition in Anko-

jarinja, In this case, a “+-" only is recorded.

M44, and M+ size

Large M4 and M4 talonid are related

characters and more common among siruc-

turally ancestral didelphines than dasyurids,

Very few dasyurids have the M4 talonid as

relatively large as it is in Ankotarinja, The M4

lalonid of most dasyurids is laterally e¢om-

pressed with one or at most two cusps present,

ihe hypoconid and entoconid or hypoconid

and hypoconulid, In this character, Ankorarinja

is most similar (among dasyurids) to Neo-

pRascogale and to a lesser extent Phascoloso-

rex. Three distinct cusps may sometimes be

present on the M4 talonid of other dasyurids

such as Murexia, Myoictis Gray, 1858 and

some Antechinus but in these forms the M4

trigonid is relatively larger than in Amkovarinja,

while the talonid is transversely compressed,

Relatively large M4 talonids characterize many

didelphine groups. For example, the talonid of

M4 in Marmosa is only slightly more reduced

than that of Ankotarinja. However, in Mar-

moya the whole M4 is not as reduced relative

to Mg as it is in <Anketarinja. Philander

Brisson, 1762 (33460), and Metachirus Bur-

meister, 1854 (J3461) also show a relatively

unreduced M4 talonid. Coona and Guggen-

heimia Paula Couto, 1952 have M4 talonids

even wider than the trigonids. Extremely

narrow My talonids (comparable with the con-

dition in most dasyurids) are found among

didelphines in species of Minusculodelphis and

and Marmosopsis,

Overall reduction of M4 relative to M4, such

as occurs in Ankararinja, does not occur in

any modern dasyurids. It is common only to

some didelphoids. M4 of Ankotarinja is un-

known but maxillary fragment UCR, 15343 in-

dicates that this tooth was as wide but not as

long antero-posteriorly as M+ of most didel-

phids. It was comparable in length to Mi of

some modern dasyurids such as Neophaseo-

gale, but wider than that tooth in most modern

dasyurids, Size of M4 in modern dasyurids

Fig. 3. A-E, scanning electron microscope photographs of Ankotarinja tirarensis. A~D, UCR, 15308,

LM}. .4, stereophotographs. E, UCR, 15343, maxillary fragment with alveoli for LM3-&

Fig. 4. A-E. scanning electron microscope photographs of Ankotarinja tiraréensis. A~C, F7332, LM3. A.

stereophotographs. D-E, Holotype, P18190, RMa-j, entoconid broken off RMj.

Fig. 5. A-E, scanning electron microscope photographs of Ankorarinja tirarensis. A-B, UCR, 15340,

dentary fragment with LM$-4. D, UCR, 15341, dentary fragment with LMé-, E, F7334, dentary

fragment with LMs-4.

62 M. ARCHER

appears to be related to relative length of the

cheek-tooth row, being shortee and more

reduced in forms with more compressed tooth

rows. This compression commonly occurs in

more sirictly carnivorous forms where

elnphasis is on development of metacrista-para-

cristid shearing elements, In more insectivorous

forms, paracristaanetacristid shearing elements

are relatively less reduced, resulting in a reda-

tively more functional M% paracrista anc

larger M4 talonid,

Reasons for overall (eduction of M4 relative

ta Mi afe not easy to interpret, In some dip-

rotodont marsupials where reduction of M4 is

advanced, sometimes even involving Insx, Py

is developed as a lage sectorial or even

plaviaulacoid tooth, possibly reflecting a shift

anterjorly m shearing emphasis, reducing the

importance of M4, In Anketarinja, although

P4 is large, the size is not particularly different

frem that of some didelphines which lack

reduced M%,

My, trigonid width, paracenid reduction, and

size of P&

Reduction of the M4 paraconid and Py are

related chsracters and often accompanied by

transverse compression of the Mj trigonid und

increase in relative importance of the proto-

conid. The relatively Jarge P4{ and wide tri-

gonid of M{ in Ankorarinja are unmatched

among living dasyurids, Even in Neopheasco-

gale the My triganid is transversely compressed

with gross reduction of the paraconid. In

Ankotarinja the M4 paraconid is very low on

the trigonid but not strongly deflected out of

alignment with whe other lingual cusps such as

occurs in most dasyurids, The closest match

among dasyurids is found it species of Smin-

thopsis, Murexla, and Keeuna but even here,

the M4 paraconid ts shifted anreriorly relative

to that cusp in Ankofarinja. A relatively wide

trigonid on M( occurs in most didelphoids, and

in part, reflects the relatively large Py m most

of these forms. In dasyurids which show pre-

molar reduction, it is P_ which ts reduced or

jost in the lower dentition (including the in-

teresting case of Planigale gifesi Aitken, 1972,

Archer 1976a), while in didelphoids and bor-

hyaenids it is Pi) which is normally reduced.

The only marked exception are specics of

Zygolester Ameghino, 1898, which are unique

among didelphids in having an extremely re-

duced, although two-rooted, P4. Reis (L957)

notes that in the M4 the paraconid is more

huceal in postin than is normal for the group.

This modificution is much tess than that seen

jv the trigonid of dasyurids with a comparably

reduced P{. Reduction of Pi normally occurs

among dasyurids which exhibit a compressed

cheek-tooth row, increased carnassiulily as

judged by proportionate in¢rease im metacrista-

paracristid letigth, enlarged canines, ete.

(Archer 1976b). Attendant reduction of the

M4 paraconid and increase in size of the Mj;

Protoconid shift the premolariform-molarifurm

boundary posteriorly. The M4 trigonid func-

fions as a stabbing, piercing premolac rather

than a sectortal tnigonid. Fucther, in dasyurids

which have lost P4, such as Daxyayrus Geallroy,

1796 and Sarcophilas, the metacrista-paracris-

tid length of M& is greater than thal of Mi, or

My. The net effect is to cancentrate the sec-

torial Function of the molars at a pom pos.

tevior of the middle of the molar row. Posterior

shift of the premolariform-mofanform boun-

dary may be scen in this way as merely main-

lainiog the structurally ancestral relationship

between these two types of teeth,

Axkorarinja is clearly structurally ancestral

in this regard and more similar te didelphids

than dasyurids.

Position of the cristid obliqua

The Mé cristid obliqua of Antotarinja

lirarensiy is unlike almost all dasyurids in that

it intersects the trigonid at a potnt so far buccal

to the metacristid carnassial notch. This con-

dition is approached in Neephuscogale and ta

a lesser extent in Keeuna, while in other

dasyurids the cristid obliqua tends to intersect

the trigonid just buccal to the carnissial swtch.

Th some dasyurids (e.g. Neophascogule and

Dasyurus) the protocenid fiank cantnhules to

the cristid obliqua on M4-4, and this same con-

dition occurs in Ankotarinja, at least on Mi,

Clemens (1966) suggests that lower teeth

(except perhaps dP4) of pedionryils can he

distinguished from species of Alphadon in thal

the crista obliqua in pediomyids intersects. the

base of the trigonid well buccal to the meta-

erishid fissure, This pediomyid condition is alsi

Present in all modern didelphids examined in

the present study and appears to be present in

itustrations of species of many of the Sopth

American fossil didelphines. The condition im

Ankorarinja is closer to this pediomyid and

didelphine condition than itis to most modern

dasyurids or Alphadon.

Althoueh the functional significance of this

difference is not clear, a relatively more buccul

posilion produces a larger tulonid basin. Posi-

lion Of the cristid Obligu|| must aiso reflect, posi-

tion of the paracone, a more buccal position

MIOCENE MARSUPICARNIVORES 63

indicating a relatively reduced or more bticcaliy

situated paracone.

The condilion fouwsd in Ankorarinja and

some dasyurids (c.g. Daryurvs) of a small

anléfiot component of the cristid obliqua

formed by the protocanid flank results in the

development of a amall accessory carnassial

notch agalnst which shears the paracone. This

makes an effective point-cutting unit that

supplements thse developed on the trizonid,

Transverse metacrintid

In Ankoterinja the metacristids sre trans.

Verse to the long axis of the dentary, In

dasyunds, this condilion ws present only in

species Uf Siinthopals, Aniechinoemys Kreift,

1867, and fo a lesser extent, species of Ningaui

and some species of Planigale. In other

dasyunds, the metaconid is displaced pos-

teriorly relative to the proloconid resulting in

the metacristid and paracristid formimg a more

obtuse angle. The tranaverse condition is

present among some but not all modern didel-

phines, some Cretaceous didclphoids, and many

Tertiary didelphids,

Among dasyurids, markedly non-transyerse

metacristids occur only in the more carnivorous

forms. This structural trend is noted by Bens-

ley (1903) who regards it as modification

towards longitudinal and away from transverse

shear, In this respect, the lower molars of

Ankotarinja demonstrate the structurally primi-

tive insectivorous condition, which is more

common among didelphids.

Stylar cusp size and position

Terminology of the stylac cusps of Anko-

tarinja used here is set out elsewhere (Archer

1976h).

The stylar cusp condition in Ankotarlnja ts

closet to that of didelphids than dasvurids in

having & large stylar cusp posterior to stB, an-

terior to stD and buccal to the low point in the

pata-metacrista of M3, which is the homologue

of the normal didelphid and variably present,

but invariably small, dasyurid stC, Further,

stD in Ankotarinja is smaller and slightly more

posterior in position than that cusp in modern

dasyurids. These non-dasyurid-ike features are

common among Cretaceous didclphines such as

glasbiines, some species of Alphadon anil

Pediomys.

Stylar cusps do not have occlusal counter-

parts in lower molars, Vet they clearly sustain

wear. This wear must result from food abrasion

during inilial puncturing prior to the cutting or

shearing occlusal phase. As the dentary closes,

forte is apphed to food trapped belWeen teeth

by the lower molars in opposition fo the whole

of the upper molars including the stylar shelf.

The area of the crushing or piticluring surfave

is increased by Jarger stylar cusps. In marsupi-

carnivores, stylar cusp reduction occurs tn the

more camivorous forms such as Sarcoplilis.

Thylacinus Temminck, 1824. and borhyaenids

where perhaps the puncturing value of these

cusps is overshadowed by the feed to have

large and sturdy shearme crests, The stvfnr

cusp size and arrangement in Ankoturinjia may

therefore be interpreted as evidence for insecti-

Vorous rather than carnivorous habits.

Metaconule development

In Ankorarinja the metaconule of the upper

molars is 7 prominent feature, while the proto-

conule is not present in M4. Conule develop-

ment is present in some dasyurids and many

didelphids. It is well-developed in most Creta

ceaus didelphines where both protoconules and

metaconules occur. Simpson (1928) notes that

these cusps in recent didelphids are represented

at most by vestiges.

The possible functional significance of meta-

conules is discussed elsewhere (Archer 1971}.

In addition, well-developed metaconules may

serve as shearing counterparts for the hypo-

conid and buccal edge of the hypocristid,

Sumunery

The dentition of Ankorarinja resembles that

of many didelphids, and some dasyurids such

as Neopharcogale, Murexia, and Sminthupsis.

Because of the middle Miocene age of the

deposit, as well as the fact that two relatively

more modern-type dasyurids (Keeune and an

unnamed form noted by Stirton, Tedford, &

Miller 1961) are in the same deposit, it would

be absurd to regard Ankorarinja as the ances-

tral dasyurid. However, it does preserve charac-

ters which could be regarded as structurally an-

cestral lo modern dasyurids. It t& clear that

many South American Tertiary didelphines of

Paleocene age (Graham & Ride 1967) share

characters with Ankotarinja which it docs not

share with moder dysyurids. Taking into

account all available morphological characters

jozether with what is currently known of their

distribution, and recognizing that the charac-

ters (incisor number and dP4 morphology)

which will distinguish between the two lincages

Dasyuroidea and Didelphoidea wee not pre-

served, one cannot avoid concluding that the

data availuble indicate a marsupicarnivore-

prohably betonging to the Dasyuroidea, but

which, like the slightly older phalangeroid

Hynyardia Spencer, 1900 (Ride 1964b), also

Fig, 6, Specimens of Keeuna woodburnei and their measurements (mm). 4, Holotype, PI8!91, RMS.

B, F7333, maxillary fragment with LM2 (broken), LM3 and alveoli for LM$. C, UCR, 15346,

dentary fragment with LP4 erupting and alveoli for LC\-M4. D, F7334, LM4. £, UCR, 15344,

LM: or Mé, F, tentatively referred to K. woadburnei, UCR, 15347, premaxitlary fragment, with

alveoli for RIt-4: d represents diastema between alveoli for REY and RI?; pmx.v. represents

edge of premaxillary vacuity.

retains a number of characters of a sort which

reyeals its derivation from early Tertiary didel-

phoids or the didelphoid-dasyurid stem. If it

should be discovered that Ankotarinja possesses

a didelphotd incisor number, the author ‘will

have no hesitation in describing it as a didel-

phoid with characters foreshadowing dasyurids,

but it is clear that it is far too late in time to

be an ancestor to the whole dasyurid family.

For the present, it is listed as ?7Dasyuridae.

Genus KEEUNA nov.

Type species; Keeuna woodburnei sp. nov. (by

designation and monotypy)-

Generic diagnosis: Differs from other Austra-

lian and New Guinean dasyurids including

Ankotarinja in combining, relatively reduced

stD on M2-%, Jarge M4, antero-posteriorly

shortened M*, and virtually absent posterior

cingulum on M4-4 (although mild posterior

cingular swelling present on holotype), and

relatively unreduced P4.

Origin of generic natne> Kee, central Austra-

lian Aboriginal word for wild cat; una, central

Australian Aboriginal word for forgotten

(Smith 1880). Keeuna is here given masculine

gender,

Keeuna woodburnei sp. nov.

FIGS 6-8

Holotype: P18191, isolated RM%.

Type locality; Tedford locality, Etadunna For-

mation, Lake Palankarinna, Etadunna Station,

S.A. (28°47'S, 138°25E).

Diagnosis: That of genus. Relative develop-

ment of entoconids, stylar cusps, and size may

prove to be diagnostic species characters.

Origin of specific name; The specific name is

in honour of Dr M. O. Woodburne who helped

MIOCENE MARSUPICARNIVOKES 65

find all of the malerial referred to this species

and who, with Dr W, A. Clemens, gave the

author fs first opportunity to study Australian

Tertiary fossils.

Referred specimens: UCR, 15271, RMit:

17333, left maxillary fragment with M® and

part of M3, UCR, 15347, right premaxillary

fraginent; UCR, 15344, isoluted LM 4; UCR,

13348, left dentary fragment; UCR, 15286, tri-

gonid RM 4; UCR, 15269, trigonid RM 4;

UCR, 15274. isolated Me; UCR, 15345, Icfi

dentary fragment; UCR, 15346, dentary frag-

ment with LP4 erupting, F7334, isolated LM4,

Description

Maxillary fragment (F7333) has LM%, pos-

lecior part LM, and alveoli for LMt. M4

narower than M3 but relatively hittle reduced

anlero-posteriorly, being only slightly shortcr

than M4. Metacone toot M4 large, equivalent!

in size to protacone root, and only just smaller

than paracone root. Interdental fenestrae occur

between M2 and M3 and between M% and Mt.

Maxillary root of zygoniatic arch arises buccal

to M*-4, Infraorbital canal opens on anterior

edge of maxillary fragment, dorsal to anterior

end M&,

Premaxillary fragment (UCR, 15347) may rep-

resent K. woodhurned on basis of size, because

larger than would be expected for other known

Negapakaldi forms. Four alveoli present.

Largest represents RL}, (alveolus broken}. This

alveolus separated from alveolus for R1I® by

Very short diastema, Alveoli for RIS-4 contact

one another. RE alveolus inclined posteriorly

and suggests RI} inclined antero-ventrally.

Based on alveolar size, RI1 largest incisor,

RIS exceeded RI4 in Jength which exceeded

RI4 in length. This may be misleading sinve

posterior lohe of RI4 (occurrence of which is

not uncommon among modern dasyurids}] may

have caused this tooth to be longer than R12.

Also, root for RI} commonly large in modern

dasyurids, while crown muy be very reduced.

Posterior to RI! alveolus, premaxillary wall

descends into pit which is ovclusal counterpart

of RC. indicating RC4 large and caniniform.

M5 with continuous anterior cingulum con-

necting parastylar corner of tooth to preproto-

crista (piece of enamel missing from anterior

cingulum of holotype). Posterior cingulum

absent (F7333) to doubtfally present

(PIS191) as swelling at base of crown abave

metacone root. Swelling not continuous with

Postprotocrista whereas this is the case in

moder dasyurids whth undoubted posterior

cingula, Five distinct buccal cusps present, StA

Gecurs at buccal end of anterior cingulum,

hetween parastylar commer of tooth and stB.

Almost vertical, minor crest copnects stA to

parastylar corner of tooth, that part of tooth

which would overlap postero-buccal edge ol

P+, Very minor, more gently inclined crest

comects sLA to stB. StH connected to paracone

by long paracrista. Buccal crest extends pos-

teriorly From stB to contact small, jow stC

which is adjacent to another small low stylar

cusp of uncertuin homology. These two stylar

cusps not connected hy crest, The posterior

small cusp connects to larger stD by Minne,

inclined crest. StD largest stylar cusp, bul

stialler than (fat cusp in M% of modern

dasyurids. StD connects lo metastylur corner

of tooth by lung, low crest, No evidence of

stE. From metastylar corner of tonth, three

crests radiate: buecal crest to stD. metacrista

fo Metacone; and minor short crest that extends

amero-lingually from metastylar cover and

ends within short distance of toothy corner.

Crescentic enamel ridge occurs lingual to mic

point of ectoloph, and buceal to mid-point of

para-metacrista, Ridge may represent cusp

analogous (o similar structure in some speci-

mens of Srinthepsis virginlaeg Tarragon, 1847.

Paracone shorter than metacane, and two

cusps widely separated. Small but clear meta.

conular jidge extends from base of metacone

to middle of postprotocrista. Postprotocrista

terminales as steep-sided crest adjavent to base

of metacone. Deep pyramid-shaped fossa

exists between hases of paracone and metacone

and buccal flank of protocone. Metucrista

approximately 1.5 {imes length paracrista.

E¢tofiexus in ectoloph slight, poimt of inflectinn

immediately posterior to siC,

Mé@ with continous antenor cingultim, Pos-

terior cingulum absent. At least four srylar

cusps present. Parastylar extension of tooth

small and accordingly litthe overlap of imeta-

stvlar cornet of M3 occurs, SiC connected to

StB as in M2, On anterior flank of cusp called

stC (F7333), small, possibly distinct swelling

occurs which may be homologous with stC of

M5. If so, cusp posterior tn that cusp in MS

might be homologous with single conspicuous

cusp in this position on M3. Posterior to stC

of M&% ts slightly larger stD. Posterior to stD,

and connected to it by crest, is stE, This cusp

exlends postertorly es ridge which terminates

short of metastylar corner of tooth. Only meta-

crista radiates from metastylar corner of tooth,

Lingual Io stC, as in M5, crescentic ndge

occurs Which may be distinct cvsp. In addition

66 M, ARCHER

TABLE 2

= Pa]

2 2 % =

a Se ¢ = s s & & i 8

S = §s§ = & S&S F€ ze | € SF = € F

= = 3 = = & = € 3 = 3. = 3 = 3

= = S S = 3 — 4 = = = 3 G =

Unusual charagrers: >: = 3 € S &- 8&8 $ § & § F = F

of Koenna *# 5 & = fi = Sc S& = B® FE Be

1, Large Ms i, i i i a i is cs ell

2, M* lack post, cing, | + | Fo ~ $$ +t + — —| = + + +

4 ArmallsiD tek + S Ft ff $F HF KE KE HK —— +

4. Compresacd M+) — Se a ee — —» om aa ae

to metaconular ridgc, as in M2, clear imeta-

conular swelling present on pestprotecrista, Na

clear proteconnle er protoconular ridge

present. Metucrisin less (han 1.5 times length

Paracrista, Ectoflexus M*# broad and relatively

deep. Point of inflection in ectoflexus occurs

anterior to stC. Otherwise morphology of MS

as in M3.

Dentary with two branches of inferior den-

lal canal, one emerging at point Tevel with

middle of M4 on buccal surface of dentary

while other emerges at point beneath posterior

root Py (latter condition determined fram

juvenile dentary, UCR, 15346). UCR, 15344

only specimen with premolar alveoli preserved.

Pi, erupting in this specimen. As result.

apparently crowded premolar condilion may

besome less crowded in adult dentary. Pre-

molar gradicnt suggested by alveoli; Py shorter

than P4 which js subequal in Jength to P4,

DP, alveoli suggest tooth as large aus P4{ and

Iwo-rooted, P4 alveoli acutely oblique with an-

terior root postero-buccal to C4 alveolus. Pos-

terior root Pi immediately posterior to C4

alveolus. Py and «P4 alveoli suggest iccth only

mildly out of alignment (although crown

alignment need not be reflected in root align-

ment), C4 alveolus relatively small, suggesting

tooth no wider than Pa.

Py partly erupted snd partly obscured. Tooth

single-cusped with tall protocontd. Paracristid

steep, No anterior cingulum cusp. Metucristid

more gently inclined and appears to direcily

contact very small posterior cingulum cusp. No

buccal of lingual cingula evident.

M; talonid wider than trigonid, and trigonid

more laterally compressed than What structure

in| Ankerarinja, Anterior cingulum relatively

well-developed, terminating lingually with

hypoconulid notch, and terminating buccally

anterior to buccal base of protoconid. Posterior

cingulum slightly shorter than anterior cingu-

lum and extends to contact hypeconulid. Pro-

Hounked crmgulum development, as bulac

hetween bases of protocanid and hypocenid,

No lingual cingulum: Paraconid low on crown,

approximately same hetght ax entoconid.. Proto-

conid tallest trigonid cusp, Metaconid just

shorter than prntocomid, Hypoconid subeqial

to entaconid in height. Paracristid complete

hetween protocanid and paraconid, and

anterior part of crest steeply inclined. Meta-

cristid and apparently paracristid fissures

extend below cutting edges of crests. Crest

descends from posterior wall of melaconid and

meets crest from anterior wall of entoconid.

‘Two efests meet with shallow, open fissure,

Crest development from posterior wall of cnto-

conid slight to absent. No crest links entoconid

and hypoconulid. Hypocristid extends postera

lingually from hypoconid to hypoconulid. Both

hypocristid and metacristid clearly not trans-

verse to long axis of tooth, Cristid obliqua in-

tersects trigonid well buccal to point below

metacristid fissure:

Isolated molars posterior ta M4 not been

posilively identified. UCR, 15274 and UCR,

15344 tentatively regarded as tepresenting M4

and M‘ respectively, This conclusion based an

trigonid width and paraconid height. Both in-

crease posteriorly in most modern dasyurids

between M4 and Mé.

UCR, 15274, LM%, trigonid just narrower

than talonid. Hypoconulid notch between paru-

stylid and lingual end of anterior cingulum

larger than in M4. Cingulum development

hetween base of protoconid and hypoconid

does not produce buccal convexity in crown

olitline. Paracomid higher on crown than in

M+. Paracristid fissure well-developed below

crest. Cristied obliqua intersects trigonid ans

extends short way up trigonid wall at point

lingual te paint of same intersection in M4.

Otherwise morphology UCR, 15274 same as

Mj.

Ger, 15344, LM4, trigonid and talonid sub-

equal in width. Hypoconulid natch larger than

in M4 but subequal to that of M4, Posterior

MIOCENE MARSUPICARNIVORES 67

cingulum Jess convex posterlorly than in M4 ar

Mi, Also, posterior cingulum extends lingually

ad contacts hypoconulid rather than stopping

short of it as in M4 and Ma, Basal cingulum

between protoconid and hypaconid well-

developed but does not cause buccal convexity.

Crests linking metaconid and entoconid Jess-

developed than in M4 and Mg (although M4

damaged in this region). Cristid obliqua inter-

sects trigonid as in Ms. Otherwise morphology

Mg, a3 in Mg.

Discussion aud comparison

A summary of important dental characters

in marsupicarnivores in general is given else-

where (Archer 19766) and to avoid repetition,

discussion of dental characters of Keeuna is

largely restricted to those characters which

eillher make Keevna unusual among the

Dasyuridae or sugeest aMnities outside of that

family. Comparison within the Marsupicarni-

vora is restricted to the Dasynridse and Didel-

phidae. because no other families contain Forms

even remotely similar to Keeuna,

Keeuna cannot be referred conclusively to

the Dasyuridae for the same reasons given

above in the discussion of Anketarinja. How-

ever, the features of Keeuna@ are more suigges-

tive of known dasyurids than didelphids, If the

referred premaxilla does in fact represent

Keeuna, there is no reason to doubt its

reference to the Dasyuridac, This specimen

shows alveoli for four incisors, The diastema

behind the anterior alveolus confirms that ihe

anterior alveolus represents 11. No dasyurid or

didelphid known to the author has a diastema

within the incisor row posterior to 13 or an-

terior to 15, other than a variably preseni dia-

stema between U4 and 13.

Although the dental characters of Keeuna

are all present in one dasyarid or another, con-

sidered together: 1, the large M4 (inferred

from alveoli): 2, virlual Jack of a posterior

cingulum on the upper molars; 3, relatively re-

duced stD on M%; and 4, antern-posteriorly

shortened M4; they make Keenna unique

among dasyurids. These characters are com-

pared in Table 2 for didelphids and dasyurids

which provide the closest similarities in upper

molar morphology to Keeuna. Many South

American Tertiary didelphids which have a

lower molar morphology (ez, Adtrandatherium,

Paula Cotto 1962, fig. 4) simifar to that of

Keeuna, are unrepresented by upper molars

and not included in Table 2. Characters 1-4

are the same as noted above. A “+" records a

condition similar to or closely apptoaching thal

found in Keeuna, A “? indicates elther thal

the dentition is too incomplete or poorly pre-

served to enable determination. A “—" indh-

cates a condition unlike that im Keeund. Poly-

lypic genera which have some forms similar to

but othets differing from Keenna are recorded

only as “sl. M* indicates any of all upper

molars.

M4 size

The possible significance of a large M4 is

discussed above. Keewra exhibits similarities in

this regard la many didelphids but only a few

struclurally ancestral dasyurlds, meluding

Atkoterinja.

Postenlor citigula on upper molars

Elsewhere (Archer (1976b) i has been

noted that cingula have the effect of increasing

molar surface area. [i is alsa possible that pos-

terior cingula on upper molars act as supple-

Mmentary shearing crests in opposition to the

paraccistids which come into effect after the

paracristids shear past the metacrista. Distribu-

tion of this character in modern dasyurids

(Archer 19766) does not appear to lend itself

to phylogenetic interpretation, being present in

some but not other species of single genera

such as Anilechinuy, Absence in Keeuna,

although perhaps phylogenctlically unimpor-

tant, is useful in combination with other

characiers for differentiating the gents.

Sylar cusp D

Small size of this cusp allres Keeura with

Ankotarinja, a5 well as. with many didelphids.

The possible significance of stylar cusp size

1s discussed. above in regard to Ankorarinja.

Compressed M4

M# of Keeuna is longitudinally compressed

tu comparison with structurally ancestral

dasyurids such as Neephascegale hut not in

comparison with structurally derived dasyurids

such as Seniatiepyis whose molars are even

mote compressed, Bensley (1903) liotes reta-

tive compression in some dasyurids and regards

this as a more insectivorous thau carnivorous

adaptation. Extremes of non-compression, such

aus occur in Sarcoppilus, result in shearing

crests which approach longitudinal rather than

a transverse crientalion,

Detaled comparisons

Overall, Keevna more closely rescmbles

some dasyunids tocluding Phascolosorex, Neo-

phascogale, some Antechinus, Murexia, and

Ankotarinja, than it does didelphids. Particnlar

similarities and differences are noted below

68 ML.

Similarittes which extend to all dasyurids are

not noted.

Phascoloserex; Similaritics include relatively

large M+ and metaconule. Upper molars of

Keeuna differ from those of Phasecolosarex

(e.g. AMNH, 109758, 151992, 101975 and

LO9757) in that stylar cusps mutch closer to

buceal edge of crown; small stylar cusp occurs

lingual io st© (although in some specimens of

species of Phascolaseres such as AMNH,

151992, this Cusp suggested on M5); M*

shorter antero-posteriorly; paracnsta ond metu-

cTista enclose more acute angle; and Mt-+ lack

clear postenor cingula, In lower molars of

Keewna, trigonid and paraconid of M{ much

less reduced; crests From posterior face of

metaconids much better-developed; talonid M:,

relatively slightly wider.

Neophayeopule: Large M+ of Keeuna similar

to that of Neophascogale (tg, AMNH,

109524), Differences in upper molars of Keewra

include these noted above in comparison with

Phascolesorex as well as lack of distinct antero-

lingual low crest developed on base of prote-

cone (Which has nothing to do with preproto-

crista); ectoloph and para-mctacnsta relatively

more widely separite at their closest point: pro-

toconule slightly better-developed. Lower

molars Keenna differ in having less reduced

Mj trigonid: relatively shorter, wider molars;

cristid obliqua Which intersects trigonid in rela-

tively more lingual position (notable in M4~-4)5

lack of post-entoconid crest which directly con-

nects to bypoconulid: relatively lower talanids,

higher trigonids; relatively shorter talonids,

Anteckinuy: Resemblance with some Anie-

ghinus (e.g. A, mayert (Rothschild & Dollman,

1930), AMNH, 109816, A, sp. AMNE,

150877 Erom New Guineas, and A. trelanure

(Thomas, 1399), WAM, M5517) considerable

including overall propornions of M%-8: some-

what similar reduction of st1D on M®*: relatively

unreduced P4. Upper molars Keeune differ in

that siD relutively slightly more reduced on

M3; stB relatively more posterior on ectoloph:

all stylar cusps relatively smaller; M4-3 some-

what sharter antero-postertorly; M4 notably

longer in proportion to length of M4; posterior

cingwla absent; metacone and paracone M4-+

ARCHER

relatively closer in height; larger, more veon-

spicuoua cusp of crest occurs lingual to stC;

ectoflexus in relatively more posterior position;

protocone shorter antero-posteriorly at its

longest poinf, Lower molars Keewee differ in

having less-compressed trigonid on M4 with

larger puracond; relatively wider talonid on

Mz; lack complete buccal cingulum such as

occurs on Mé~, of some Antechinus species

(eg, A. mayeri); lower molars relatively

shorter, wider; enfoconids M4-5_ relatively

taller; hypoconulid wider and extends farther

from: postero-lingual comer of M4-4.

Murexia: Similarities between ‘species of

Murexia (eg. M. fongicaudata (Schlegel

1866) (AMNH, 101972 and 152035)) in-

chide comparable relative length of M4; rela-

tively unreduced P4. Upper molats of Keewria

differ in same features from Murexia as they

do from molars of Antechinux except us

follows, In Keeway all stylar cusps except C

relatively smaller; posterior cingulum of upper

molars virtually undeveloped (although only

slight postertur cingular development occurs in

species of Murexia); M$ relatively longer;

metaconular crest from base of metacone

less well-developed and Iscks low, minor crest

linking stD with metacone (latter observed

only in unwom Specimens of Murexia

examined in this study, AMNH, 152035).

Lower molars of Keeuna differ in same features

from teeth of Murexia that differentiate teeth

of Antechines, except as follows. In lower

molars of Keeyna entoconids relatively shorter

antcro-posteriorly, and higher; buccal cingulum

absent (occurs in one specimen of Mureala.

AMNH, 152035); low direct crest linking pes-

terior face of entoconid with hypoconulid

absent.

Ankotarinja: Comparison with much smaller

Ankatarinja demonstrates that bath farms simi-

lar tn having relatively small stylar cusps (par-

ticularly stD); atylar cusp(s}) present betweed

B and D; no posterior cingulum; complete an-

terior cingulum; slightly smaller paracones than

melacones; Lick of direct crest linking ento-

conid with hypoconulid; relatively large P4; un-

reduced trigonid and paraconid of M{- Upper

molars of Keeuna differ from those of A4nke-

farinja in larger stze and more gntetior position

Fig. 7. A-F, scanning electron microscope photographs of Keenan woodburnei, A, Holotype, P1811,

KM2, stereo photographs. B, tentatively referred lo K. woodburnei, UCR, 15347, prensaxillary

fragment with alveoli tor RLY-5 and posterior edge of alveolus for RI, C, F7334, LM4. D, UCR,

15344, LM or LM&, E, UCR, 15346, dentury fragment with unerupied LP4 aad alveoli for

1.P4-M% and edges of alveoli for LO{ and LMé,

jaa]

Ss)

—

jaa

jaa}

}

MIOCENE MARSUPICARNIVORES 7!

of stD on M3-3; number of cusps in position

of stC; presence of siylar crest limgual to siC;

less. well-developed metaconule,; presence of

deep pit hetween bases of paracone und meta-

cone, Lower molars of Keene differ in having

longer metacrista; relalively larger M4; more

lingual intersection of cristid obliqua and in-

Zonid; non-transverse inetacristid and hypocris-

tid; large hypoconulid of Mg; antero-posteriurly

non-compressed inigonids; relatively unequal

heights of the paraconids and metaconids. Dif-

ferences in position of intersection of cristid

obliqua and trigonid in <Ankoturinja and

Keeune not one of kind, but degree, position

being telalively more lingual in Keeunes,

Other comparisons; No other dasyurids warrant

dctailed comparisons. Most didelphids reveal

fewer similarities, particularly if the referred

premaxillary fragment of Keeuna does in fact

belong to this form. General similarities with

Cretaceous didelphines (e.g, some species of

Alphaden such as A. rhelster) include ihe

paracone, which although smoller than the

metacone, 1s not markedly so. Other similarities

between Keeung and species of Alpfiddon it-

clude relatively reduced atylar cusps (except

8), particularly D on M& of species such as 4.

rhoissery and A. Jullf Clemens, 1966, and

presence uf slylar cusp or crest lingual to sic

in A, rhaister. Marked differences in upper

molars of Keeuna include much smaller proto-

ahd metaconules; less-deeply concave and sym-

metric ectoloph of M4; stnaller stB; and absent

postenor cingulum (present in some species of

Alphadon). In lower molars, similarities

between Keryna and species of Alphadon in-

clude relatively unreduced ingonid of M4, Dif-

ferences in lower molars of Keexra include

much smaller paraconid relative to metaconid

in Ms-3 a8 opposed to specimens referred to

species of Alphadon (Clemens [966, Lille-

graven 1969),

Compited with holarctic Tertiary didel-

phines, relative size of paracane in Keetina is

similar to condition seen in species of Pera-

dectes and Peratherident. Other similarities in-

clude relatively fow stylar cusps; unpeduced

condition of Py (and presumably P+) and M4

paraconid; lack ul posterior cingulum on upper

molars; relatively shallow indentation of ecto-

loph; extreme buccal position of styfar cusps:

and evidently oon-iransverse onentation of

metucristid and hypocristid. Differences in

Keeund include relatively larger stylar cusps

(particulurly D): relatively poorly-developed

metaconule; non-pediomyid—ike cristid obliqua

Urientation; relatively taller metqconid and en-

toconid, and shorter paraconid: and presence of

rudimentary buccal cingulum between bases of

protoconid and hypoconid, Despite these dif-

ferences, Keeuna more eloscly resembles these

uidelphines than any others for which good

illustrations or photographs are available.

Close similarities may exist between Keeuna

and some Paleocene (Riochican) didelphids.

Untortunately very few are Enown from upper

teeth and few are adequately illustrated. As

noled ubove, all modern aidelphines examined,

and mast Tertiary didelphines exhibit a pediv-

mytid type of cristid obliqua, which diflers from

Keeuna. Of all Paleocene forms ilfustrated hy

Paula Couto (1952, 1962, L970) and Simpson

(1947), Mérandatherium igs pevhaps most like

Keeuna, Keeuna differs from this didelphid in

felatively smaller size of Py and presence of

buccal cingulum developed between bases of

protoconid and hypoconid. Upper molars of

Mirandatherium are unknown,

Other didelphids do not reveal enough simi

laritics to warrant separate comparisons,

Summary

Teeth of Keeuna, although resemblmeg teeth

of some Tertiary didelphines such as species of

Peratherium, Peradectes, and Mirandatherinwn,

are broadly similar to teeth of some anodern

dasyurids such as New Guinean Antechinus, In

view also of the Australian localiry of Keene

it seems most logical to regard this form as a

somewhat unusual dasyurid, probably withour

direct descendants in the modem dasyurid

fauna. Resemblances between upper molars of

Ankotarinja and Keeuna further suggest the

possibility thal these two forms are more

closely related to each other than either is to

other dasyurids, thereby placing Keeuwra in 3

structurally intermediate position between

Ankotarinja and made dasyurids,

Discussion of Etadunna marsupicarnivores

Stitton, Tedford, & Miller (1961, p, 35)

briefly describe {but not name) another carni-

Vore fram the Etadunna Formation. According

to their description “The size of the animal is

comparable to Dasyrrus quoll .. . The three

Pig. 8. 4-C, scanning ¢lecttun microscope photographs of Keetna woodburmel, F7333, maxillary frag-

Ment with partial L.M3, LM and alveoli for LM: C, stereo photographs.

72 M, ARCHER

premolars with gradation in size from P4 to Py,

and (he absence of the metaconid on M4 sug

gests that this animal may not be far removed

from the ancestry of Thylacinas’. The author

has seen drawings and photographs of this

Specimen (courtesy Dr W. A. Clemens and

Mr C. Campbell) and it is clear that nothing

else about the specimen, including the mor-

phology ef the upper molars, shows any simi-

larity 10 Thylactnus. On the contrary, it appears

to represent another dasyurid lineage (perhaps

related 16 Dyttyerus) in which metaconid re-

duction his occurred only on M4, This un-

named dasyurid and Keexte ate the only Eta-

diinna camiveres which can be referred with

some confidence to the Dasyuridae,

Ankeravinia ts either a didelphid or dasyurid.

Compared with known didelphoids, dasytraids

and perameloids, the preserved portions of

Anketarinja do not enable confident reference

to a particular marsupial family. It is referred

to here as ?Dusyuridae.

Ankeotarinja and Keéuna compared with

modern dasyurids, share most Uenial chanw-

fers with living New Guinean species of Nea-

phascegale, Phescolosorex, and Antechinus.

Similarities with Australian forms are fewer

and those that do exist are with forms found

in generally non-arid Australian habitats, Broacl

My, talonid, large M4, aartrow and relatively

uncrowded premolar tow, and Jarge entoconids

are characters either lacking or sare in Aus-

tralian arid-adapted dasyurids, Several authors

(c.g. Woodburne 1967, Schodde & Calaby

1972, Stirton, Tedford, & Woodburne | 96%)

indicate that New Guinea hag many mammals

(eg. species of Dendrolagus Muller, 1839,

Doreopsis Schlegel & Mulles, 1839, Diy-

teechuris Peters, 1874, Microperorycies Stein,

1932. Murexia, Myoictis, Neophaycogale and

Phascalosorex) from highland rainforest habi-

tats which appear to be structurally ancestral

within their respective families.

Evidence for a Iless-arikl central Australia

during Ngapakaldi time is reviewed by Stirton,

Tedford, & Woodburne (1968). In addition,

pollen from the base of che Etudunna Forma-

tion, recently obtained from bores in South

Australia, has been found to include Notho-

fagus sp. (pers. comm., W, &. Harris, South

Australian Department of Mines), a genus of

plants presently restricted in the Australian

fegion to the high-rainfall habitats of eastern

Australia, New Guinea and New Zeulanct,

Resemblance between Negapakaldi marsupicar-

hivores and Iilying New Gumean dasyurids,

suggests these living highland New Guinean

dasyurids may have avoided verlain selective

pressures brought to bear on mursupicurnivores

living in central Australia, following Ngapa-

kaldj time. These pressures may have included

progressive deterioration of climate with in-

creasing aridiwly. Elsewhere, as part of revision

of the dasyurid genus Sminthopsis, the author

has given evidence for believing that several

Australian dasyurid lineages underwent arid:

adaptation, Arid-adapted forms now dominate

the majority of Australia and are in clear con-

trast to the mursupicarnivores of the Npapa-

kaldi local fauna deseribed here.

Acknowledgments

] am deeply indebted to Drs W. A. Clemens

and M,. 0. Woodbume of the University of

California at Berkeley and Riverside respec-

lively, for giving me the opportunity to help

collect and study Australian Tertiary Fossils jn

1971, Dr Woodburne wis-also a field associate

and co-worker in 1972, These trips were

financed by a National Science Foundation

Grant to Drs Woodburne and Clemens, Dr

R. H. Tedford, American Musewn of Naturul

History, kindly allowed me to examine in the

field dasyurid and other fossil material

recovered from deposits in the Lake Frome

Basin, South Australia, in 1973, Aspects of the

1973 trip were financed by a» National Science

Foundation Grant to Dr Tedford. Other people

wha helped make the 1972 collections included

Messrs R, Lawson, W, Head, South Australian

Museum, and my wife Elizabeth. Mr C.

Campbell of the Universiry of California at

Berkeley allowed me to examine photographs

of dasyurid material he a at present examining.

Dr Clemens provided casts of upper molars of

Lance species of Alphudon. Drs W..D- L. Ride,

Western Australian Museum, and A, Bartholo-

mui, Qucensland Museum, read and construs-

lively critized drafts of this manuscript. Mrs C.

Farlow and Miss P. Rainbird of the Queens-

land Museum typed drafts of the manuscript.

Messrs A. Easton and A. Elliot of the Queens-

land Museum helped with ospects of light

photography. Mr J. Hardy, University of

Queensland, produced the excellent scanning

cleviron microscope photographs_

MIGCENE MARSUPICARNIVORES 73

References

ArcHer, M. (1974)—The developanent of the

Cheek-teeth in Awicehinus flavipes (Mar-

supialia, Dusyyridae), J. R, Soc, West Aust.

87, 54-43.

Arcoer, M. (19750},—The developtnent of pre-

molar and molar crowns of Avitechinus

flavipes (Marsupialia, Dusyuridae) and the

significance of cusp ontogeny in mammalian

tecth. J. R. Soe. West, Awst. 57, 118-325-

Arcues, M. (1975h).—ingamt, a new genus of

tiny dasyurids (marsupialia) and two new

species, trom atid Western Austrslia, WN.

dimealey! and N, ride’. Men, Gd Meas, 17,

27-29.

Arcuen, M. (1975c).—Abnormal dental develop-

ment and its significance in dasyurids and

other marsuplals, Afem, Gd Mrs. 17, 251-265.

Archer, M, ¢1976a).—Kevision of the niarsuplal

dasyucid genus Plenigele Troughton, Ment.

Od Murs., in press.

Arcurk, M, (1976b).—The dasyurid dentition

and its relationships to that of didelphids,

thylacinids, borhyaenids (Marsupicarnivora)

and peramelids (Peramelina). Avs, J, Zool.,

Suppl. Series, in press.

Bensvey, H, A, (1903).—On the evolution of the

Australian Marsupialia; with remarks on the

relatiouships of the marsupials jn general.

Trane. Linn, Sav. Lond, (Zeal.) 9, 83-217,

Clemens, W. A. (1966).—Fossil mammule of the

type Lance Formation Wyoming Part I.

bres aiele. Bull, Dep. Geol, Univ, Calif, 62,

1-122,

Cremens, W. A. (1975).—Fossi) mammals of the

Type Lance Formation Wyoming Part_ TIL.

Eutheria and summary. Aull. Dep. Geel.

Univ, Cali7, 94. 1-1u2,

Counins, L. R. (1974) —"“Monotremes and mar:

supials.” ¢Smithsonian: Washington, }

Guava, $8. F.. & Ripe, W. DBD. L. £1967) —Intra-

class Metatheria. #7) “The Fossil Record”.

(Geol. Soc. Lond: London.)

Laure, B. M, o.. & Ant, J, B, (1954)>—"List of

land mammals of New Guinea, Celebes and

adjacent islands !758-}952." (Tonbridge

Printers: Tonbridge. }

Littrcraven, fF. A. (1969),.—Lalest Cretaceous

mammale of upper part of Edmonton Forma-

tion of Alberta. Canada, and review of mar-

supial-placentzl dichotomy in mammalian

evolution. Paleent. Conir. Cuiv, Kans. Article

50 (Vertebrata 12), 1-122.

Pauays Coura, ©, bE (1952).—Fossil mammals

from the beginning of the Cenozoie in Brasil,

Marsupialja: Didelphidac. Aimer. Mus. Novit.

1567, 1-26,

Pau, Coura, C, ne | 1962)—Didelfideos fossiles

del Paleoveno de Brasil. Cienciay Zophigious

%, 135-166.

PauLa Coupo, C. pe (1970)—News on the fossil

marsupials from the Riochicun of Brazil. Ad.

stead. brast!, Clee. 42, 19-34.

Ree ©. A. (1957) —Sobre la posicion sistemuticu

ile "Zyyulestes paraneisis” Amegh, y de

“f“ysaless erirérrianus” Amegh. con una

reconsidetacion de Ja edad y cortelucion det

"Mesupotamiense”. Molmberzice 5, 209-226,

Rive, W. D. L. 11964) —Anlechinis rosamendae,

4 new species of dasyurid marsupial rom the

Pilbura District of Western Australia: with

remarks on the classification of A tiechinas.

Ww, Aast. Nat. 9, 58-65.

Riot, W. D. L. (1970) —"A eulde to the pative

manimals. of Australia.” (Oxford Univ. Pr:

Melbourne.)

Rostinson, R. (1966)—‘Abariginal myths and

legends,” (Sun Books: Melbourne.)

ScitoppE, R,, & Canasy, J. H. (1972) —The bio-

geogrophy of the Australo-Papuan bird and

mamnial faunas co relation to Torres Strvit,

In D. Walker (Fu.) “Bridge and barrier: the

natural and cultural history of Torres Strait?

{Ast Nat. Univ, publ, B6/3¢1972); Can-

berra,)

Stor, B. (19721.—La faunnvle de mammiferes du

Crétucé supérieur da Laguno Unsayo (Andes

péruviennes). Ball. Mus, natn, Uist, mat. (3)

99, 375-405.

Simpson, G. G. (1928),—American Eocene didel:

phids. Auner. Mus. Novit. 307, 1-7-

Simesow, G. G. (1929).—American Mesozoic

+ dig Mem. Peabody Mus, Yale 3, 1-

Stupson, GG. (1941)—The affinities of ihe Bor-

hyaenidue. Amer. Mfus. Navit. 118, 1-6.

Simpson, G, G, ¢1945) —The principles of classi-

fication and a classification of Mamitals. Rall,

Amer, Mus. nat, Mist, 85, iexvi, 1-339.

Simpson, G. G. (1947)—A new Eocene mar

supial from Brazil. Amer. Mus. Navit, 357,

1-7.

Smeson, G, G. (1948)—The beginalne of the

age of mammals in South America Part J. In-

troduction, Systematics: Marsupialia, Eden

tala, Condylarthra, Litopterna, and Notiopro-

vonia, Bull. Amer, Mus. nat, Fisr 94, 1-232.

Sineiam, W. J, (1906)-—Mammalia of the Santa

Cruz ‘beds. Marsupialia, Rep. Princeton

Exped. Patagonia 4, 333-460.

Suri, J. (1880), —"The Booandik Tribe of South

Australian Aborigines: «a sketch of their

habits, cusiome. legends, and language”

(Gove. Printer: Adelaide?

Smror, Ro A., Trpeorp, ROW. & Mitirr, A. A.

(£961)—Cenozoie stratigraphy and verte-

brute paleontology. of the Tirari Desert, South

Australia, Rec, S. Aust. Mus, t4, 19-61,

Stimtox, R. A. Teprorn, Ro A. & Woonpvenr,

M. QG. (1968),—Australian Tertiary deposits

containing letrestrial mammals, Bali, Dep

Geal. Unite, Calif, 77, P30.

Tuomas, ©. (1887)—On the homologies and

sucession of the teeth tn the Dasyuoridae

Phil. Troms. R. Sav, 178, 443-462.

Weooorcrne, M, ©, (1967).—The Alcodte Fuuna,

Central Australia: an integrated palcontalogi

cal and geological study. Bur. Min. Resor

Aust. Bull, 87, 1-187.

Weons, H. E. (1924),—The position of the

“sparussodonts'! with notes an the relation

ships and history of the Marsupialia. Mull,

Avior Maz Nut. Hist. $177,101.

SOUTHERN AUSTRALIAN SPECIES OF CHAMPIA AND CHYLOCLADIA

(RHODYMENIALES: RHODOPHYTA)

BY D. J. REEDMAN* AND H. B. S. WOMERSLEY

Summary

REEDMAN, D. J., & WOMERSLEY, H. B. S. (1976).-Southern Australian species of Champia

and Chylocladia (Rhodymeniales: Rhodophyta). Trans. R. Soc. S$. Aust. 100 (2), 75-104,

31 May 1976.

Five species of Champia are recognised on southern Australian coasts. C. viridis C. Ag. (including

C. tasmanica Harvey, C. oppositifolia J.Ag., and C. verticillata J.Ag.) resembles the type species

C. lumbricalis (L.) Desvaux from South Africa] in having numerous scattered longitudinal

filaments passing through the diaphragms of the thallus. C. insignis Lucas from Tasmania also has

scattered longitudinal filaments. Three other species, C. affinis (Hook. & Harv.) J. Ag. (including C.

obsolete. Harvey), C. zostericola Harvey (including var. arcuata Hook. & Harv. of C. affinis) and

C. parvula (C.Ag.) Harvey var. amphibolis var. noy., have only peripheral longitudinal filaments in

the thallus. Australian records of the South African C. compressa Harvey probably apply to juvenile

C. tasmanica or to a probably undescribed species on the N.S.W. coast.

One species of Chylocladia, C. grandis, is newly described. Other Australian species previously

referred to Chylocladia belong to Lomentaria or are relegated to synonyms.

SOUTHERN AUSTRALIAN SPECIES OF CHAMPIA AND CHYLOCLADIA

(RHODYMENIALES:; RHODOPHYTA)

by D. J. REEDMAN* and H. B. S$, WomersLey*

Summary

ReepmMan, D. J., & Womerstey, H. B. $. (1976).—Southern Australian species of Chanipia

and Chylocladia (Rhodymeniales: Rhodophyta). Trans. R. Soc, S. Aust, 100(2), 75-104,

31 May 1976.

Five species of Champia are recognised on southern Australian coasts. C. viridis C. Ag,

tincluding C, tasmanica Harvey, C. oppeasitifolia J,Ag,, and C, verticillaia J.Ag.) resembles

the lype species [C., lumbricalis (L.) Desvaux from South Africa] in having numerous scattered

longitudinal filaments passing through the diaphragms of the thallus. C, iasignis Lucas. from

Tasmania also has scattered longitudinal filaments. Three other species, C. affinis (Hook. &

Harv.) J. Ag. (including C. obsoleta Harvey), C. zostericela Harvey (including var, arewata

Hook. & Harv. of €. affinis) and C. parvula (C.Ag.) Harvey var. amphibolis vur. nov.

have only peripheral longitudinal filaments in the thallus. Australian records of the South

African C. contpressa Harvey probably apply to juvenile C. tesmanica or to a probably

undescribed species on the N.S.W. coast.

One species of Chylocladia, C. grandis, is newly described, Other Australian species pre-

viously teferzed to Chylocladia belong t Lomentaria or ate relegated to synonyms,

Totroduction

Champia Desvyaux is in general a well

characterised genus (Kylin 1956) of the family

Champiaceae Kuetzing (1843) (syn, Lomen-

tarinceae Naegeli 1847) of the Rhodymeniales

and includes ntimerous species from most

cousts of the world. Some 9 specics have been

credited to southern Australia, and ss with so

many Australian genera, taxonomic distinctions

ure uncertain and in some cases the carlicst

Valid names are not in current use.

Ciylocladia Greville has been credited with

several Australian species, all of which are

synonyms of species of Chanipia or other

genera. One new species of Ciiylocladia has,

however, been found in South Australian

waters,

The type species of Champia is C, luambri-

calis (L.) Desvaux (1808, p. 246), from the

Cape of Good Hope, South Africa, Although a

well-marked species, C. /uinbricalis has never

been studied in detail, and the generic concepts

of structure and reproduction are based largely

on the European C, parvula (C. Agardh) Har-

vey which has been investiguted most recently

by Bliding (1928) who reviews earlier studies.

The thallus construction and reproduction of

the type species do, however, appear to con-

form with those of C, parvula, and a brief

account of the type species is given below.

Champia is characterised by a multiaxial,

hollow but septate and thus segmented, usually

much-branched thallus which originates from

a ring of apical cells (or a ring plus several

central cells). These apical cells each cut off a

filament of cells which runs longitudinally

through the thallus, and from the peripheral

filaments the continuous cortical layer of cells

originates close to the apex. If a group of

central apical cells is present (as in the type

species), then longitudinal filaments also occur

in the central region throughout the “hollow”

thallus. The characteristic transverse dis-

phragms in the thallus are derived from the

Iongitudinal filament cells very close to the

apex, and originate either from alternate cells

or ones 2-3 cells apatt. Each longitudinal fili-

ment cell cuts off cells laterally in one trans-

verse plane, and these link up and divide fur-

ther to form the characteristic 1 cell thick dia-

phragm, the peripheral filaments. being adja-

cent to the cortex or sometimes separated by

* Department of Botuny, University of Adelaide, Adelaide, S. Aust 5000_

16 QD. J, REEDMAN & H, B.S. WOMERSLEY

one diaphraym cell. The longitudinal filament

cells lying between the diaphragms usually cut

off one (-3) gland cells, Outer cortical layers

may be formed, and in some species an inner

cortex ul chizoidal filaments develops,

Most species are much branched, and the

branches atise from the regions of the dia-

Phragms by development of 4 Ling of apical

cells (rom cortical cells of the parent branch.

Reproductively Champia is fairly distinctive,

The procarp consists of a +-celled carpogonial

branch borne on a supporting cell fa cortical

cell), together with a 2-celled auxiliary cell

branch, present before fertilisation. The cysto-

carps ure external and sub-spherical to urceo-

late. asiiolate, containing a carposporephyte

with 4 basal fusion cell and much branehed

gommablas! filaments with terminal carpo-

sporangia. The inner cells of the pericarp

become stellate and form a network (“tela

arachinoidea").

Spermatangia are cut off from mother cells

Uerived from the witter cortical cells, and may

cover extensive jreas of the bratiches.

Tetrasporangia develop by enlargement of

inner cortical vells. they are tetrahedrally

divided ynd vecur scattered over the branches.

Species of Chapin are moderately common

along most of the coast of southern Australia,

und sre commonly mentioned in ecologicul

uccounts, In general, however, they do not

characterise any communities, though C. affinis

may be common in shallow water om rock plat-

torms (Womersley 1948, p, 158),

Chylocladia Greville differs from Chrmpia

in that the cystucurps do not have an ostiole

and the carposporophyte consists of a large,

basal fusion celi giving rise directly ta carpo-

sporangia. The thallus construction of the two

genera ts similar.

CHAMPIA Desvaux

The strycture and reproduction of the type

species, Chamipia lumbricalls

Champia lanbericaliy (L.) Desvaux (1808,

p. 246) is bused on Ulva luiibeicaliy Linnaeus

(1771. p. 300) from the Cape of Good Hope.

The species is a distinct one, and material from

Camps Bay, Cape Vown, South Africa (CG.

Dieckmann, FLvilit973) has been studied to

check On ihe generic characteristics:

Cy lumbeicalig is a robust species forming

clumips with numerous axes to 15 ent high,

arising Cran an entangled, stoloniferous base.

The axes are 2-4 mm thick. tercte and linear,

with ovcasional basally constricied branches

with rounded apices. The diaphvagms are regu-

lar in position, $-1¢~1£) mim apart and largely

obscured below by the thick cortex.

The apices have a central group of 7-8 ini-

jials and peripheral ring of about 14 Imitials,

resiiting in Jengitudinal filaments passing

(hrough the inner part of each diaphragm as

well as 14-2!) wround the periphery. One com-

plete (rarely 2) and two part longitudinal fila-

ment cells occur between the diaphragms. bear-

jing |-3 gland cells. The corlex close to the

apex becomes 3.4 cells thick, with a dense

outer cortical layer of anticlmally clongate

cells. In old axes the cortex increnses to about

8 colls thick, and o weft of rhizoids also deve-

Jops as an internal layer’ to the cortex.

The reproductive organs occur on tufts of

short, adventitious, branchlets (5-10 mm long

und 1-1+ mm thick) formed on the upper hail

of the axes. Usually the tufts are dense, with

numerous, curved {concave acduxially) tertile

branchlets, but in some plaints cystocarps are

borne on single lateral branchlets,

Cystocarps are borne mainly on the adawial

(concave) side of the curved branchlets, uflen

with 2-4 grouped together, they are ovoid with

a small ostiole, i-2 mm in diameter ancl high.

The carpospurophyte arises from 4 basal fusion

cell, with a much-branched gonimohlast bear-

ing terminal carposporangia; subterminal cells

probably also mature into carposporangia after

Joss of the terminal ones. The inver cells of the

pericarp become stellate, forming a loose

tissue. and the outer wall is comparutively

thick. Curpogonial branches and early post-

fertilisation stages have not been observed.

Spermatia form a continuous covering all

around the branchlets, or sometimes luryely on

the adaxial sides, with the outer cortical cells

culling olf 2-4 elongate spermatangial mother

cells which then cut off several avoid spermu-

7aTVeaL,

Teteasporangia occur densely in the brunch-

leis and are transformed from most of the large

inner comical cells; they are slightly pyrifarm

to avoid, abeut 100. 2m long, and tetrahedrally

divided.

The above description of C. lumbricalix

agrees Well in cssential generic details wih that

of Bliding (1928, p. 5) for C. parvude. thuugh

the latter is much smaller find slendgrer, wilh &

much thinner cortex, and bus only penpheral

longitudinal filaments: ‘Uhere js thus ne reason

to doubt the generic concept of Champa as

secoyiised by Blidge and hy Rylin (1956, 7

346),

SOUTHERN AUSTRALIAN CHAMPIA AND CHYLOCLADLA vt

Key to southern Australian species of Champia

1, Vhallus with longitudinal filaments scattered

thoouzh the diaphragms as well as pevipheral

filaments; branches usually lincar, Msally con-

stiicted or not, not or slightly constricted al the

cisphragms; branching irregular or distichous

Thallus with peripheral longitudinal filaments

enly; branches usually tapering to base and

apex. usuilly slightly to moderately constrieted

at the diuphragms; branching irregular or aise

2, Branching irrewular, often distant, branches

lieews, 14-) t-2! mm browd ......,

C, yirielis (p. 77)

2, Branching subdistichous, fairly regularly pia-

nale; branches 2-5 mm broad, with a hitsul

stender stalk ae, Cotnsizets bp B14

Thallus segments obscured; hooked branches

absent; Usually epilithic; cortical eclls cutung off,

near branch apices, Usually several small outer

cortical ces wt first around their margins, liter

becoming almost continuous over the thallus as

an outer layer and in older parts cutting off

further outer cortical cells; inner primary cor-

ticn] cells, which thus become obscured, are

ovoid, 14-2 times as lang as broad. and 20-30

ant broad oo... C. affinis (p. 82)

3. Thallus segments clearly defined throughout!

most of thallus; usually epiphytic on seagrasses

or larger algae; cortical cells each cuting off

usually only 1(-2) small cells fram their corners,

so that the single Jayer of large cortical cells

remains clearly defined (hroughoul most of the

thallus: cortical cells usually ungulur, 2-3(-4)

times as long as broad _ A

4. Branchlets 4-( mm. branches 1-2 mm, in

diameter; cortex essentially single layered

throughout; usually one compleie longitudinal

filament cell between diaphragms; ultimate

branchlets often hooked; usually epiphytic on

Posidonia, Amplibolis or Sarger algac

C. zastericola (p. 87)

4, Granchlets $/3-4% mm, branches 4-1 mm, in

diameters cores mostly single layered but 2-4

cells thick tm oldest axes; usually two. com-

plete longitudinal filament celly between dia-

phraems: ultimate branchlets usually linear.

rarely hooked; epiphytic on Applibalis -

Cc, paevula var. amphibatis (p. 91)

Champia viridis C, Agardh 1828: 115. Kuet-

zing 1849; 842,

hmimiatt viridis (C, Agardh) Trevisan [848:

Las.

Champia tasmanice Harvey 1844a: 407, pl 19,

847: 78: be59. 307, J. Agardb 1852: 370,

1876; 206; 1879: 67, pl. 19, fis 10-12, De

Toni 190Ua; 79; 1800h: 563. Guiler 1952) 95,

Hooker & Harvey 1847; 402. King ef al., 1971;

122(%), Kuetzing 1549: Bél: 1868: 30, pl S4e.

Kylin (930: 79. Luvas 1909» 34; 1929n: $9;

1929b; 50 Lucas & Perrin 1947: 207. fig. 72,

May 19645: 362, Okamura 1904: 88, Reinhold

1897: 53; 1899:> 45. Shepherd & Womersley

1970: 1345 1971; 165, Sonder 1846: 177(7);

1853; @82> 1855: S518; 1880, 17, ‘Late 1kR2!

18. ‘Tisdull 1893) 506. Wilson 1892: JO.

Womersley 1950: 176; 196fir ISE.

Champia tasmanica var. gracilis Harvey 1863;

synop.; 27, Sonder 1880: 17 Tate 1&b2: 1K.

enidaldia tasmanica (Harvey) Trevisan 1848

108.

Champta onpositfolte J. Agarih 1901) 27. De

Toni 1924= 309, Kylin 1931: 29, pl. 16, fig. 47.

May 1965: 362,

Champta verticitlaw t Agardh 901; 24, De

Yoni 1924: 309. Kylig 1931: 29, pl 17. fig 39.

May, 1965; 362.

Champla compressa sensu Harvey 1843, synop,:

27 tat least in part—see below),

HGS |, 24-D, 10

Thalias (Fig. 10) usually with several main

axes from a. stoloniferous base, forming .a

dense, spreading tuft commonly 5-15¢-20) cm

high, moderately or slightly adhering to paper,

medium to dark tek er red-purple in colour.

Axes (1-12-3 min broad. usually with numer:

ous irregular branches in 24 orders, often sub-

Opposite, usually less than 1 cm (sometimes 2-4

em) apart, in older or grazed plants ofien ver-

licillately branched. All branches terete to

slightly compressed, (!-)1-24 mm_ broad,

linear to slightly curved. basally constricted

and with rounded upices. Diaphragms 4-1(-14)

nium spart, regular and usually conspicuous in

surface view of thallus. Corre of a single layer

of compact cells, polygonal and 25-40 pm

across in surface view, with an inner cortex of

rhizoidal filaments in older parts of robust

plants. Longitdinal filaments both peripheral

and central, with one complete and two pan

filament cells between each diaphragin,

Cystocarps scattered over young branches,

globular to urceolate, ostiolate, $- 1 mm int ia

meter. Apparently very few cystecarpic plants

have been collected.

Spermatangia forming a continuous layer

over beanchlets.

Terrasporangia scaliered in younk branches,

60—J 20 uni in diameter.

Type lecalin, W. Aust

Type. Herh, Agardh, LD, 26112.

Divrihation. Prom, Rottnest TL. W. Aust,

around southern Australia and Tasmunia to

Gabo f.. Vic., Usually on rough-watcer coasts or

in strong currents, fran shaded pools to 25 mM

deep, with a slender form on Pesidonia in more

sheltered waters,

The Lype specimen of C. viridis C, Ayardh

consists of 8 small branches on mica, und ts

identical with the later described Cy tasnanice

Harvey (type from Tasmania, n TCD), under

which name mast specimens of this taxon have

been known.

Fig.

DB. J. REEDMAN & H. B, 8. WOMERSLEY

gop OOD a,

750 um

Champia viridis. A, Longitudinal section of a branch apex showing the development of cortex,

longitudinal filaments and diaphragms (A42995), 8. Surface view of a branch apex showing

central and peripheral apical cells (A42991), C. Cross section of a young branch showing @

diaphragm with peripheral and scattered longitudi

mal filaments (A42991). D. Three dimen-

sional view of thallus showing diaphragms and longitudinal filaments with gland cells

(A42991). E, Longitudinal section of un older axis showing development of rhizoids from the

peripheral diaphragm cells (A3055()),

SOUTHERN AUSTRALIAN CHAMPIA AND CHYLOCLADIA 79

Harvey (1863, synop. p. 27) described var,

gracilis of C, tasmaities, A stritable lectolype

is prabably Harvey's, Alg. Aust. Exsicc. 251,

tn TED, fran Port Phillip, Vic, and speci-

mens in MEL (45227) from Brighton. Port

Phillip, Vie. (Harvey, Trav. Set 483), named

var. fracta, are probably the same, These are

small, slender forms. often found on Posidonia

under moderate conditions of water movement

(e.g. Port Phillip, Vic., St Vincent Gulf, S.

Aust.) in carly summer, and dowbtfully justify

a yarietal name. The thallus is usually 4-10 cm

high, densely tufted with an entangled base,

irregularly branched, with occasional curved

branch apices, branches mostly 4-1 mm. broad

and segments #-1 times as fong as broad,

slightly constricted at the diaphragms. The cell

structure is very similar to larger forms typical

of the species, with cells about 40 pm broad,

(1-)2-3 times as long as broad, angular with

small cells cut off from the corners (about as

many small cells as parent cells). While ex-

tremes of this shelreted-water form and the

robust rough-water form appear relatively dis-

tinct, a good range of intergrades does occur.

C. verticilleta JT. Apardh (L901, p, 26) is

bused on a specimen (type in LD, 26078) from

Port Elliot, S. Aust. (Hussey) and ts an older

plant of C. viridis with verticillate branching,

and C. oppositifolia J. Agardh (1901, p. 27),

with type in LD (26148) ts « plant with some-

what more distinct opposite branching.

C. viridiy 1s a distinctive species in s7e,

form, and in having central as well as peri-

pheral longitudinal filaments. It varies con-

siderably im robustness and thickness of

branches and tn branching, with Irequent

occurrence of prolifcrous branches giving a

subverticillate arrangement. These variations

are cilher ecological ar duc ta the age of the

plant.

C. viridis js most closely related to the type

species. C. lumbricalis; from South Africa, but

Abbreviations used in Figures 1-9.

is distinct in being a less robust plant and pot

developing a cortex several cells thick,

A Preiss specimen in MEL (45206) is C,

zostericola, not C_ viridis, bat a small form of

the Intter does occur in Western Australta.

C. viridis (as C. taymanica) has been

recorded fram New Zealand by Naylor (1954,

p. 658), This recerd has nor been checked, but

may apply to the clasely related but distinct C.

novae-zelandiae Huoker & Hatvey, which has

central longitudinal filaments but a many

layered cortex,

STRUCTURE AND REPRODUCTION

Material studied: Nora Creina, 5. Aust, upper-

most sublittoral (Reedman, 12.31.1973; ADU,

A42995); Cape Lannes. 5. Aust., under ledges

(Reedman, 10,i1,1973; ADU A42991), and Staple-

ton Point, Prosser Bay, Tas., 8-12 m deep (Olsen,

71.¥i.1966; ADU, A30S50),

Thalluy development. The apex of a branch

(Fig. 14, B) includes both a central group of

12-16 initisls and a peripheral ring of initials,

producing longitidina! filaments passing

through the central as well as the peripheral

regiona of the diaphragms (Fig. 1€, D), as in

the type species. The number of central longi-

tudinal filaments is commonly greater than the

number of apical initials, due apparently to

division of the initials and consequent branch-

ing of the filaments and subsequent loss of

some initials and terrmnation of some filaments

at diaphragms, The apical initials of the peri-

pheral ring divide transversely, and within 2

or 3 cells of the apex divide periclinally form-

ing cortical cells which then divide..anticlinally

to form the single-layered cortex (Fig. LA).

Alternate cells of the longitudinal filaments cut

off cells laterally which join to form the single-

fayered diaphragms (Fig, 14), after which the

tongitudinal filament cells become very elon-

gate, The alternate cells of the longitudinal fila-

ments, lying between the diaphragms, form

1(=2) gland cells (Fig, 1). Only the cortical

tell formed directly from the longitudinal fila-

a — apical cell feb — fused carpogonial 0 — asliole

acl, — apical groupe branch per — pericarp

omc —-atuxiliary mother cel) = Fu — fusion cell re — thizoldal cell

uux — autiliary cell f1, 22 — gonimoblas! cells Sp = Speropatangnint

ch —curpogonialbranch ~— ot) — yonimolobe Pr spérwataogiel mother

ou outer cortical cell ald — gland cell aii — supporting cell

cort = — inner carlical cell | — longitudinal filament 1 — tela arachnoides

csp = — carposporangium le — tateral connecting uu — trichogyne

d — diaphragm ocll Ispp = — Letrasporangium

AG D. J. REEDMAN & H. B. S. WOMERSLEY

& cs

B®

0 08

oe)

ee Sg

Q80n &

ce Paty ag -

*~ Ba RD ETS

CSEOF ik

Fig. 2. Champia viridis, A. An early post-fertilisation stage showing supporting cell with auxiliary cel]

branch and carpogonial branch with fusions occurring between the cells. Considerable cortical

proliferation has occurred towards formation of the pericarp (A30550). B. A post-fertilisation

sige showing a connection between the fused carpogonial branch and the auxiliary cell. and

formation of the first gonimoblast initial (A30550). C. Development of tetrasporangial initials

{A30550). D. A mature tetrasporangium (A30850). Champia insignis. E. Surface view of a

branch apex showing initials (A12237). FL Section of male thallus showing development of

spermatangia (A12237).

mient cell is in pit-connection with it, In older

parts, s0me cortical cells may produce small

outer cells from their corners, hut the cortex

Temains essentially only one cell thick.

In older branches, rhizoidal cells develop

from peripheral diaphragm cells and form a

loose layer lining the inner side of the cortex

(Fig. 1D), as in the type species.

Branches arise from the region of the dia-

phragms, following development of u group of

apical initials from the cortical cells.

Procarp and carposporephyte

Cystocarpic specimens appear to be rare, und

only one female specimen with very young

carposporophytes has been available. The

carpogonial branch is 4-celled und borne on a

SOUTHERN AUSTRALIAN CITAMPIA AND CHYLOCLADIA Da)

supporting cell which also bears the auxiliary

mother cell with its auxiliary cell (Fig. 24).

Following fertilisation, the cells of the carpo-

gonial branch fuse and a connection between

the fused carpogonial branch ancl the auxiliary

cell is formed (Fig. 28), and the first gonimo-

blast cell ig cut off from the auxiliary cell,

Early post-fertilisation development is accom-

panied by division of the surrounding cortical

cells to form the pericarp (Fig. 243, and in

the One specimen (Stenhouse Bay, S. Aust,

7 m deep, Kraft, 18.tx.1973; ADU, Ad4564)

observed with cystocarps, the structure of the

pericarp and carposporophyte is typical of

Champia and very similar to the illustrations

of Bliding (1928) for © parvula.

Spermatangia

Spermatangial plants have pot been ob-

served.

Tetraxporartgia

Tetrasporangia develop hy enlargement of

cortical cells (Fig. 2€, 2) and are tetra-

hedrally divided, 60-120 jam in thameter when

mature, and scattered in younger brauches.

NOTES ON CHAMPIA COMPRESSA

Champia compres\a Harvey IS38: 402;

1847: 78, pl. 30) was first recorded from Aus-

tralia by Harvey (1863, synop,: 27) on the

basis of specimens from Western Australia

(Clifren, and Alg, Aust. Exsice, 250A) and

from Port Fairy, Vie. (Alg. Aust. Exsiec.

250D). Since then, C. cenipressa has been

recorded from Australia by the following

authors, probably largely on the basis of Har-

vev's records: Gatnet 1971: 96. Lucas 1909;

34, Lucas & Perrin (947: 206. Muy 1947;

275; 1965: 362. Sonder 1880; 17(2). Tisdall

1898: S506.

However, comparison with material of

Champia compressa from St James, Simons-

lown. S. Africu 0G. Dieckninn, 29.11.1973;

ADLI, Ad4601) shows that the Australian

reoords dlmast certyinly are not C. eompresse.

The South African species has strongly com-

pressest hranches with numerous central longi-

tudinal filaments scattered across the dia-

phragns. Harvey's Western Australian speci-

mens also have central longitudinal filaments

tnel the hrinches are only slightly compressed,

They agree in these features with C. viridis C.

Agardh, and are very similur lo young, well

displayed, specimens of this species Front else-

where in southern Austria, Harveys Alg.

Aust. Exsice. 250D (ti TCD) from Port Fairy,

Viz., includes C\ viridiy as well as some plants

which are not. 1 Champia, and one specimen of

250D in BM is C. zastericofa.

Other records of C. conipressa from south-

ern Australia probably apply to €. vridis if the

specimens have central bongitudinal filaments,

but the records of May (1947, p. 275, 1965,

p. 362) from N.S.W. apply to a separate spe-

cies which may be unnamed. This small, irri-

descent species, with branches attached to

others by haptera, does not have central longi-

tudinal filaments aim! is only slightly com-

pressed. It is thus distinct from both C, viridis

wnd C. campressa, and also differs from the

very. strongly flattened subtropical C. vieillirdii

Kuetzing, which from material from the Solo-

mon Islands (Womersley & Bailey 1970, p.

321) 1s so Strongly Mattened that the dia-

phragms are only 2+ cells across in the direc-

tion of flattening of the thallus, and Jongi-

tudinal filaments are almost entirely around the

periphery,

It is therefore considered that Chanzpla com-

presse does not occur on Australian coasts,

Other records of this species trom outside

South Africu also need checking; the record of

Weber van Bosse (1928, p, 477) from Borlieo

is probably C. wrefilardil, and that af Joly

(1965, p. 176) from Brazil prebably applies

to a different species.

Champia insignis Lucas 1931; 409, pl, 25. fig.

1, Guiler 1952: 94, Lucay & Perrin 1947:

207. May 1965: 362.

FIGS 2£, F, 11a

Thalls (Pig. 11.4) with one to several main

axes fo 18 ¢m high, arising from a small, slis-

coid te slightly lobed holdfast on. pebbles or

shells; branches of pyramidal form, with lower

laterals often similarly branched; thallus adher-

ing closely to paper, colour red-brown (her-

barium specimens) to “bright purple” (Lucas).

Axes 3-5 mm in diameter, subterete (possibly

slightly compressed), lincar, bearing alternate

or opposite laterals mostly $-| ci apart and

sub-distichously arranged along the ives, often

somewhat denuded towards the base. Main

jateral branches usually with a slender stalk

(Fig, 11.4), then broadening, linear or gently

lapering, usually 3-3 mm in diameter. with

rounded apex. Lesser branches similar but slen-

derer and shorter. Diaphragms apparent

throughout most of the thallus, 2-3(—+) min

apart in older parts, 1-2 mm apart in younger

branches which are slightly constricted at the

diaphragms. Cortex essentially one cell thick,

the cells subpolygonal in surface view, mostly

32 D. J, RFRDMAN & H, B, 8, WOMERSLEY

50-100 pm long and 35-50 ym broad, cutting

off 1-3 smull cells {rom theie outer corners and

more Dumerous such cells near the thallus hase.

Longitudinal filaments acattered (hroyghout the

diaphragms, consisting of several (7) cells be-

tween diaphragms,

Cywfocerps scattered over the — lesser

branches, conical to urccolate, osticlate, 2-11

mm in divnveter; catposporophyle branched

with lower sterile cells and terminal carpo-

sporangia.

Spermatangle cut off from cortical cells (Fig.

2#') and Jorming collarike patches on cither

side of the diaphragms of lesser branches.

Tetrasporangia scattered over the lesser

branches, tetrahedrally divided, 80-100 4m in

diameter,

Type locality. R, Derwent Estuary, Tas.

(“Sandy Bay, Hobart. Oct. 1925" on type

sheets, }

Lectotype. Herb, Lucas, NSW, 136559. Syn-

types (cf) im NSW (136558) and ADU

{A12237)_

Distribution. Qoly known from the type col-

lection and the following Tasmanian collec-

tions; D’Entrecastesux Channel, Nov, 1910

(NSW, 126561); Browns River, Lecus, Oct.

1923 (NSW, 136562); and Snug, Lucas, Aug.

1925 (NSW, 136560). Two specimens in the

BM, tabelled “Vas, Oldfield”, one numbered 81.

are probably alse from whe Derwent Estuary.

The species appedrs to be known only from,

or just south of, the Derwent Estuary,

Lucas did not specify type material, but the

one now selecied as jectotype is the cystocarpic

specimen illustrated by Lucas (1931, pl, 25,

fig, 13,

The above vescription is compiled from that

of Lucas (1931, p, 409) and study of the type

and other material in NSW and ADU, C. insiv-

nis bas apparently not been collected recently.

but it seems tu be a quite distinct species. It

resembles C. viridis in having peripheral and

central apical cells (Pig. 26) producing longi-

tudinal filaments scattered across the dia-

phragms, bul differs m form and dimensions

and in being essentially distichously branched.

Lucas (1931, p, 409) refers to the whole plant

as being “compressed”. It is clesirable, however,

thut Hiquid preserved collections should be stu-

died to confirm such sspects.

The BM specimens agree well with the type

cellection though the lateral branvhes do not

appear to be themselves. disttchously branched,

and the main breaches are basally constricted

but scarcely stalked; they are eystocarpic, In

calposporaphyte striigture and the ostiolate

cystocatp, C, insigniy appears to conform well

with Champia.

Champia affinis (Hooker & Harvey) J, Agardh

1876; 304. De Toni 1900a; 75, pl, 5, fig. 2;

1900b: $59; 1924: 307, Guiler 1952; 94,

Harvey 1835a: S545(7}: 1859: 307; 1863,

synop.: 27, Kylin 193]: 28, Lucas 1909: 34;

19299: 19; 1929h: $0, Reinbold 1897; 53;

1899: 45, Sonder 1980: 17. Tisdall 1898:

506. Wilson 1892: 180,

Chylocladia afftis Hooker & Harvey 1247;

402. Harvey 1847: 79, pl, 29(7),

Lonmentaria affinis (Hooker & Harvey) Kuet-

wing 1849: 863, J. Agardh 1452: 730. Sonder

1853; 693,

Gastroclonium affine (Hooker & Harvey) Kuet-

zing 1849: 866,

Eipianiadi kaliformis sensu. Harvey 1844b:

Champia ebsoleta Harvey 185%: 307; 1863,

aynop,; 27, J. Agavdh 1876: 394 De Toni

19004; 75, pl. 5, fig, 3; 1900b: 559: 1924: 307.

Guiler (952; 94, Kylin 1931: 28, pl. 1S, fig,

33, Liwas 1909: 34: 19293* 195 1929b: 49.

Lucas & Perrin 1947: 206. May 1965: 342,

Reinbok! 1898: 46. Sander 1880; 17. Wilson

1898. 506. Wamersley 1930; 176: 1966: 150.

FIGS 3, 4, 118, 12

Thallas (Pigs 11B, 124, B) erect, usually

415-40) cm high, with one to several main

axes from a smal) discoid holdfast, grey-red to

purple in colour, adhering to paper: usually

growing on rock or on Amphibolis, rarely on

Posidonia. Axes usually densely and irregularly

radially branched for 3 or 4 ofders, branches

of pyramidal form (more spreading in plants

on scagrasses), often denuded below: axes

I-3; mm, branches 4-14 mm, and lesser

branches 4-4 mm, in diameter, all branches

slightly ‘basally constricted and tapering to

rounded apices. Diaphraymy usuully fairly dis-

tinct in lesser branches, obscured on older

axes, (4-)1-14{-2) mimi apart (segments

(4—)1-14 times as long as broad}, thallus con-

stricted at diaphragms except on older axes,

Cortex of a layer of large sub-oveid cells,

(20-)25-40(-60) pm across, and a sparse

layer of outer small cells around margins af

inner cells in young branches (Fig. 37, 1), be-

coming more or less continuous on older paris

(Fig. 3G-1) and near bases of old plants 2—4

cells thick (Fig 32). Longitudinal filaments

usually confined to periphery of the dia-

phragins (Fig. 3C), rarely with 1 or 2 within

the periphery, usually with two (occasionally

Fig. 3-

SOUTHERN AUSTRALIAN CHAMPIA AND CHYLOCLADIA §3

steizes, Dery. «odie

« Ge, grrr,

Z gs wos pe hase Face ws ui hy

a cagte ack iC i ata

Kary, i

i at ! I 72! |

en oa

“AL deeocoey ye

& aa 3 ¢ ‘i

ta. ay

Os > or V

(29S SOK Sp

= ae

Champia affinis. A. Longitudinal section of a branch apex showing the development of cortex,

longftudinaf filaments and diaphragms {A42994). B, Surface view of a branch apex showing

apical cells (A42994). C. cross section of a mature branch showing a diaphragm with peri-

pheral longitudinal filaments and small outer cortical cells (442994). D, Longitudinal section

of mature branch (A42993). E. Longitudinal section of an old axis showing several layers of

cortical cells (A42997). F. Surface cell pattern of type specimen (Gunn, in BM) 10 segments

from a branch apex. G. Ditio, 30 segments from a branch apex, A, Ditto for Harvey, Ale,

Aust. Exsicc. 2521 from Georgetown, Tas., as C. ebsoleta, 30 cells from o branch apex, 7,

Ditto for A42990, 10 segments from a branch apex,

Bs 1), J) REEDMAN & H. B. & WOMERSLEY

ohe or thfec) complete cells and two part cells

herween the diaphragms,

Cysiocarps sitigle, Scattered over lesser

branches, subspherical to urceolate, 1-14 mm

long, 4-14 mm in dlumeter, with a distinel

ostiole (Fig. 4€’).

Spermatangia (Fig, 4E) in patches arentid

branchlets. on either side of diaphragms, often

envering most of the segments.

Tetrasporangia scattered over hrattches,

60-80 wm in diameter (Fig. 4F).

hectetype locality, Georgelown, Tas

(Gina).

Leclatype, BM,

Distribution, From King Geotge Sound, W.

Aust, to Western Perl, Vic, and arountl Tus-

mania. Generally found in shallow water on

reel’s on rough-water cousts, rarcly epiphytic on

Pasidonia or yobust algae.

©. affinis was reported from New Zealand

byw Harvey (18556, p, 236} and recently by

Chapman & Dromegoole (1970, p. 145). There

wre also specimens in BM from New Zealand,

but while they appear closely related ta C’

affinity there aré differences in form, Derailed

comparisons of liquid-preserved material ate

needed to establish the relationships of the New

Zeuland plant,

Hooker & Harvey (1847, p. 402) based C,

affints on plants From Georgetown, and a cysie

carpic plant in BM has been selected as lecto-

type (Fig. 118). Other specimens in BM and

in TCD are syniypes. The type specimens are

Of loose, Spreading form and typical of plants

from moderately sheltered conditions: the dia-

phragms are nol vorspicuous and the wall has

an outer layer of small eclls which becomes

cohnunawous in older parts,

Some of these Georgetown specimens are

refered by Harvey to C, ehsoleta, which he

deseribed (1859, p. 307) on the hasis of

thatlas struchury and cystacarpic plants, refer-

ting to "Alg. Exsic. n. 252" snd the follawing

localities:

“HAL, Georgetown. Southport, Cy Saart,

DISTRIB, Port Fairy. Victoria. WoW."

‘Lhe critical specimens in TCD Include

|. “Georgetown. Sept. 1848. 252 0". This speci

men (Fig. 124) was previausly considered

the lecteivpe (by H.B.S.W-. itn 1952). but is

leirasporangial and hays no name on the

sheet; it is an old plant but is C. affiniy,

being very similar to the type of this species.

Another specimen labelled “Georgetown,

V.D,. 252 1" is also ain old, baltered speci-

men, probably of C. affinis,

2. “Southport, V.D.L. C. Staart’—four speci-

mens, two with "C, obyaleta” om the sheets,

ancl which ure C, alfviis. They are all letra-

sporungtal and do not match the type des-

criplion atall well,

3. “Port Fairy, Vic. WHA, 252 D"—five speci-

mens, 3 tetrasporangial and 2 cystocarpic:

these match the description well and include

the only cystocarpic specimens jin TCD. One

letrasporangial specimen has “Cvrampia oh-

solera”’ on the sheet.

On the hasis that the leclotype spevimen. of

C. obiseleia should have Harvey's number 252

on i and should also agree well with the type

description and be cystucanpic, one of the two

Part Pairy specimens (Fi, 128) is now

selected as the loctotype and the other Port

Fairy specimens are then syntypes.

In describing C, ohgaleta, Harvey (1859, p,

307) stated “perhaps only a variety of €-.

affints’, and Kylin (1931. p. 28) doubted that

they were distinc: species. Study of Harvey's

Georgetown material, the Port Fairy specimens

of C. obseleta, and Knowledge of this commen

species along southern Australian coasts, sug.

gests strongly that only one species is involved,

and C. ohsoleta is therefore reduced to syno-

nymy. C.. affintiy occurs mainly on rocks and

platforms at about low tide level, and under

rough-water conditions in such habitats. it is

of pyramidal form and grey-purple in culour,

us Harvey (1859, p. 307) noted. While nor-

mally epilithic, it o¢casionally occurs on other

robust algae or on the seagrass Aiphibalts,

and some of Harvey's Georgetowm specimens

were growing on Posidontec, While there appear

to be no structural differences between these

forins on seagrasses and the rough-water forms,

the former are more loosely branched anid of

more spreading habit.

The type specimen of {-. affinity shows num-

erous small outer cells cut off from the primary

cortical cells (Fig, 34, @), but not as many as

in rough-waler forms (Fig. 3/). However, the

habit, luck of clearly visible primary carlicul

celly and obscuring of the diaphragms in most

of the thallus, are features of C. affints as

understood here. and differentiate this species

from C. zextericala {see helow), The type of

C. affints and other specimens (e.g. in ADU)

from Georgetown in Vasmania, Pere Phillip

Heads in Victoria. and near American River

inict on Kangaroo Island, appesr to represent

relatively galm water forms of the species,

SOUTHERN AUSTRALIAN CHAMPIA AND CHYLOCLADIA 83

and the type of C. absoleta to represent rough-

water forms of more pyramidal habit and with

more prominent outer cortical cell layers.

Some specimens of Chuampia, epiphytic on

Posidunie, resemble C. affiniy in that the seg-

ments aré nol distinct and a moderate number

of outer cortical cells are present, The segments

ure, however, offer distinctly jonger than in the

type of C. affinis, being 2—3(—4) mes as long

us bruad, The longitudinal filaments often con-

sist of 2-3 complete cells between the diar

phmgms, and the primary cortical cells are

muderately conspicuous. Most of the plants

with these charuclerislics afte stall and pos-

sibly young, though often fertile, For the pre-

sent they ate regarded as probably a form of

C. affnis, but further studies on their seasonal

growth and variation is needed. The specimens

concerned include: Port Lincoln, S. Aust,

4-12 m deep, on Posiionia {Sheplterd,

23.vill.1975; ADU, A46561, A46367, A46570),

and Pig L, American R, inlet, Kangaroo L., S.

Aust. (Womersley, 17.1,1947; ADU, A4467),

Cc. affints differs from the other common

southern Australian specics, C. zestericela, in

its habit, lack of hooked branches, and thick

2-3 lityered) cortex which obscures both the

diuphragms and the Jarge primary cortical cells,

The latter species is discussed further below,

bul very oceasonal plants with intermediate

characters do occur. Harvey's Alg, Aust.

Exsice. 253H, distributed as C. effinis, is typical

C. zostericold, and this has led to considerable

confusion,

J. Agardh (1876, p. 304) distinguished two

varieties of C, affinis (var, u affinis and var. B

intermedia) on the proportions of the articuly-

tions sod density of letrasporangia. Both these

features wre unsatisfactory characters to separ-

ate varicties, and Kylin (1931, p- 28) con-

sidered var, intermedia as intermediate between

C. affia’s and C. ebyeleta. These varieties do

hot uppear worth dislingttishing from the ape-

cits.

Hooker & Harvey (1847, p. 4021, following

description of C. «ffimis, also described var,

acvote, This varicty is considefed specifically

distinct and fs pelegated below to the synonymy

of C, 2estericola Harvey,

STRUCTURE AND REPRODUCTION

Material stidjed: Cape Linnes, S. Aust. low

eqlittoral (Reena, WGi1973; ADU, Aa2g4s

am! 127.1973; ADU, A42990); Nora Creina, §&

Aust, lower enlittornl (eed, 12,7),1973;

ADU, A42994): sud Pennington Bay, Kangaioo

1, S. Aust, low evlitioral on reef (Reedman

13.i¥,1973; ADU, A42997).

Thellus development

There ure 12-16 apical Initials (Fig. 32)

which form 4 peripherul cing of longitudinal

laments; only very oceasionally have filaments

been seen within the periphery of the dia-

phragms. Whe inittaly segment (Fig, 34) as do

the peripheral apical initials in C, weridis, bul

the cortical layer of large cells cuts off small

outer cells, at first around their outer margins,

bul a more or less continuous layer of squall

tells occurs on mature parts (Fig. 3G J), and

near the base (especially in older plants) a

corlex several ceils thick is developed (Fig.

3£)- Hairs are commonly formed from outer

corlical cells near branch apices. The dia-

phragms are fermed usually by every third or

fourth cell of the Jongitudinal filaments (Fig.

34, D), leaving usually two or three complete

cells between the diaphragms, each cell com-

monly bearing a gland cell. Rhizoidul develop-

ment within the cortical layer has aol been

observed.

Branches arise from the region of the cia-

pPowagms, where a ring of outer curtiéal Cells

becomes meristematic and forms the apical ifi-

uals of the branch. Branching cecurs irregularly

and often densely on all sides

Procarp and carposporophyte

The supporting cell (Fig. 44) is uw lorge

cortical cell in primary pil-connection with a

longitudinal filament, and is generally attached

opposite a gland cell. The eystecarps thus lie

between the diaphragms of a branch, and

cystocarpic plants are common. The support-

ine cell ix multinucleate and densely cytoplas-

mic, and cuts off a 4-celled. curved, curpo-

gonial heanch (Fig, 44), of which the first cell

is binucleate and the other three uninucleate,

The supperting cell alse produces a multi-

nugleate, densely cytoplasmic auxiliary mother

cell (Fig. 4A). which produces q unjrucleate

auniliacy cell just prior to fertilisation.

Fellowing fertilisution, the pit-connections

of the carpogonial branch cells enlarge [Fig.

48) and the cells tend to fuse. Following pre-

sumed diploidisation of the auxsliary cell, first

and then second gonimoblast cells are pro-

duced, and the latter divides further to produce

a cluster of branched zgonimollast filaments

tFip. 4C, DB). which terminate in uninucleate,

ovoid carposporangia, The catposporangia

mature simultancously, byt a new gonimolobe

sammonly develops from the basal cell of the

gouimoblasc and produces a secondary, later

maturing. amalfer cluster of carposporangia.

84 D. I, REEDMAN & H. B. 5S. WOMERSLEY

a 50 um

100 gan

50 um

Fig. 4. Champia affinis. A. Supporting cell With auxiliary mother cell and carpogonial branch

(A42990), B, Post-fertilisation stage with cells of carpogonial branch fusing (A42990). C,

Section of an immature cystocarp showing development of carposporophyte from old auxiliary

cell (A42997); D. A mature carposperophyte showing much-branched gonimoblast with ter-

minal curposporangia, surrounded by cells of the “tela arachnuidea” (A42990), E. Section of

male thallus showing development of spermatangia (A42997), F, Section with a mature tetra-

sporangium (A42993).

Vegetative cells adjacent to the auxiliary

mother cell becomes densely cytoplasmic, and

assist. nutrition of the developing carposporo-

phyte. Pit-connections between the lower ceils

of the gonimoblast enlarge considerably but the

cells do not fuse completely.

As the gonimoblast develops, vegetative cells

around its base divide to produce the pericarp.

Inner cells of this form the cell reticulum, with

aly outer wall several cells thick (Fig. 4C).

The mature cvystovarp has a well-defined

astiolo.

Sperinatangia

Spermatangial mother cells are cut off from

the small outer cortical cells to form. a con

tinuous Jayer over the branches, and each cuts

off 2-3 ovoid spermatangia (Fig. 4E). Usually

the entire spermatangium is shed.

Tetrasporangia

Tetrasporangia (Fig, 4F) develop by

enlargement of inner cortical cells which deve-

lop several secondary pit-connections with wi-

jacent cells. They are tetrahedrally divided,

SOUTHERN AUSTRALIAN CHAMP/IA AND CHYLOCLADIA 87

wilh it thick gelatinuus sheath, arnct usually

densely scattered over the branches.

Champia zostericola (Harvey) comb. nov.

Lomentaria, zostéricola Haivey 1855a;

1863, synop,: 26, J, Agardh (876: 632.

Gastrocloniunt (7) zestericolum (Harvey) De

Toni 190th: 567,

Chu teieladtat tostericoly (Aurvey) Kylin 1931:

545,

Chylocludia affinis var, arcwaia Hocker &

Harvey 1847: 402. Womersley 1966: 150.

Lomentaria affinis sensu Kueizing 1865;

86d-£.. Sonder 1855: 323,

Champia affinis sensu King et al. 1971: 122.

Lucas, & Perrin 1947: 206, fig. 71, May 1965:

362, Shepherd & Womersley IS71l; 163.

Womersley 1950: 176.

Champia parvula sensu Harvey 1855a: 548 (ia

part).

3h, pl.

FIGS 5, 6, 12€, 13

Thallus (Figs 12C, 13) usually spreading

and forming irregular clumps with several

branches from the entangled base, sometimes

with one or more erect uxes and spreading

laterals, commonly 6 to 20 cm high, grey-red

te red in colour, normally epiphytic en the sea-

grasses Pesidonta and Amphibolis or on larece

algae, possibly on rock: attachment at first by

means of a small discoid holdfast with one to

several axes, luter altaching by small adventi-

tious multicellular pads to the seagrass or itself.

Axes usually 14-3(-5) mm in diameter,

branches only slightly slenderer and ullimate

branches 4-2 mm in diameter, hranches

slightly basally constricied and with rounded

apices; Mature planis normally with some to

many branches ending in recurved ("“hooked’’)

tips (Fig, 13); young branches distinctly con-

stricted at diaphragms, segments mostly (4-}

1-14 times as long as broad. Diaphragms dis-

tinct except in aldest parts of some plants,

4—1+(—2) mm apart, Cortex single layered, of

relatively large ajgular cells [40-60(—75) pm

xeress und mostly {1/4—-)2-4 times as long as

broad] which are usually arranged more or Jess

in longitudinal rows (Fig. 5E-H). Near the

apices, each cell usually cuts off a simele, rela-

lively small cell from oear a corner, and fur-

ther such cells develop on older parts; how-

ever, the essentially single layered cortex of

large cells is maintained throughout most of

the plant (Fig. 5f, A). Longitudinal filaments

canfined to periphery of the diaphragms, rarely

with odd ones more centrally platsl, with

generally one complete cell and two part cells

between the diaphragms (Fiz. 5C),

Cysiecurps subspherical lo slightly cenical,

base hroad and slightly constricted, 2-1 mm in

diameter,

Spermatangia scattered over smaller branch-

tots, as extensive patches of collardike sore

around the diaphragms.

Tetrusporangia scattered over branches,

60-100 »m in diameter.

Type locality. Rottnest 1, W, Aust.

Type. TCD (Harvey, Tray. Set 195),

Distribution, From the Abrolhos Islands, W,

“Aust. urdund southetn Australia lo Kiama,

N.S.W. and around Tasmania. Generally epi-

phytic on seagrasses or other algae, from low

tide level ta 41 m deep, generally under slight

ta moderate water movement.

C. zostericola is based on small plants 3-1

cm high, growing on Pesidenia (not Zostera},

The type is No. 195 in Harvey's “Travelling

Set”, and his Alg. Aust. Exsicc, 294A (MEL

45197) from Fremantle (Fix [2C) ts very

similar. Harvey was in the vicinity of Fre-

mantle: from April to June 1854, and during

this period the plants are young (though often

fertile), but may not show the typical hooked

branches. Later in the year’, especially in spring

and early summer (September to November)

the planis reach 20 cm in height and nearly all

plants develop the hooked branches.

The lectorype (Fiz, 13.4) of var. arcuate

Hooker & Harvey of C, affinis has been

sclected from several specimens in BM, It ts

a well-developed specimen, attached to stems

of Heterezastera (7) and with numerous

hooked branches. Whereas the type of C.

zastericola isa young, small plant, thal of var,

arcudta is an older, larger plant of the same

&pecies,

The gencral confusion betwecn C_ affiris und

C, zostericul@ (or C. affinis var, crenata) is

probably largely due to Harvey in his Alg.

Ausl, Exsice. listing 233H from Western Port,

Vic. as C. affinis, whereas these specimens are

typical C. zosteritola.

Tn contrast to C_ affinis which is usually epi-

lithic on rough-water coasts and only occa-

sionally occurs on robust algae or on sea-

grusses, C. sestericole Is a common epiphyte

ou FPosidonie and on some targer algae,

usually in conditions of slight to moderatu

water movement and extending into deeper

water.

The presence of hooked branches, the

clearly septale rhallus almost throughout, the

essentiaily one cell thick cortex throughout the

38 DB, J. REEDMAN & H, BH, S. WOMERSITY

Fig. 5. Champia sostericala, A. Surface view of apax of branch showing apical cells (A43556), A,

Cross séciion of branch showing coytex, diaphragm and peripheral longitudinal filaments

(A43556). €. Longitudinal section of mature branch showing single layered cortex, diaphragms

and longitudinal filaments with gland cells (443556), D. Longitudinal section of an old axe

showing slight development of small outer cells (A8944). Z. Surface cell pattern of Harvey

Alg. Aust. Exsice. 249A (MFI.. 45197), (0 cells from branch apex. F, Ditto, 30 cells from a

branch apex. G, Surface cell pattern of the type specimen in BM of C. affinis. var, arcunia,

1G cells from a branch apex, H. Ditto, 30 cells from a branch apex.

plant with few small cells tying largely between

the primary ones, and the cortical cell dimen-

sions and arrangement. characterise this spe-

cles, bur occasiunal plants occur without

hooked branches. While most mature plants

have several Vague axes from their entangled

bases, some (from Tasmania and Port Phillip

Heads in particular) do have well developed

main axes with abundant laterals. Such plants

commonly (but not always) have hooked

branches and have the single-layered, large-

celled cortex typical of C zostericola. Howe

ever, the number of small cortical cells cut off

from the larger ones does vary somewhat, and

very occasionally plants intermediate in this

respect with C. affinis are found (see under

C. affinix),

C. zeslericela shows the yariation in form

which occurs in many other algal species dis-

tributed along all of southern Australia, i.e.

the western specimens are generally smaller

and less robust, and in the east, especially near

Port Phillip Heads.and in Tasmania, larger and

more robust plunts oecut.

Varition in diameter of the branches and

uXes (§ considerable, probably largely depend-

SOUTHERN AUSTRALIAN CHAMP/4 AND CHYLOCLADIA 89

ent on age, bul branches at normally over

1 mm thick, However, new growth on older

denuded branches may be slander and only

about + mm thick, as shown on several col-

lections from Pearson I., S. Aust. in ADU.

Many references to Champia affiniy apply, at

least in part, to C. zostericole rather than to

true C, affiniz (sec above), Probably most Aus-

tratian records of C, purvula also apply to

young plants of C. zostfericofa, though some

may apply to slender C. viricdiy.

STRUCTURE AND REPRODUCTION

Materia! sindied: Point Peron, W. Aust. drift

(Garden, 15,xi,1968; ADU, A34256); Port Noar-

lunga, S. Aust, 6-7 ni deep oa jetty piles aha-

son, 1941,1973; ADU, A43596); Marino, S. Aust.,

dritt (Homerstey, 26.x.1975; ADU, A46646).

Thallas development

There ate 14-20 apicil initials (Fig. 5A)

which form a peripheral ring of longitudinal

filaments (Fig, 5). with only occasionally an

odd inner one, The initials segment as in C-

offinie and a single layer of large cortical cells

ix formed, artanged more or Jess in longitudinal

lines (Fig. SE-H). Fairly near the apices, these

corticul cells become angular and cul off from

a comer a smaller cell, which remains essen-

lially in the layer of Lirger cells (Fig, SE-H}.

The smaller cells are at first simifar in number

to the larger primary cortical] cells, but later

more may be formed; however, the cortex

remains essentially only one cell thick throug-

out most of the thallus (Fig, 5C, D), The dia-

pheagms are formed usually from alternate

longitidinal filament cells, so that there is one

complete longitudinal filament (with a gland

ecll) and two part ones between successive

diaphragms (Fig. 5C), The relatively thin,

essentially single layered cortex results in the

ptimary cortical cells being visible throughout

most of the plant, and the diaphragms are a)so

conspicuous, Rhizoidal devclopment within the

cortex does not occur.

Branches arise from the region of the dia-

phragnis, with their apical cells differentiating

from the cortical cells. Near the base of en-

tangled thalli, small branches may develop into

haustorial pads of tissue and attach to other

branches or to the host,

Many of the branch apices are curved or

hooked (Figs 138, 144), and in some cases

these aid in attachment, The only structural

diflerence in hooked branch ends appears to be

that on the convex side cach sczment has 4

greater number of cells than on the concave

side,

Procerp and curposporophyte

From the limited female material available,

the supporting cell of the procarp appears to

be a large cortical cell, which produces the

Carpogonial and aéxihary cell branches (Fig,

64). Following fertilisation, the pit-connec-

lions of the carpogonial branch cells enlarge

(Fig. 68) and the cells Fuse, with a connection

forming from the old carpogonium to the auxi-

liary cells (Fig. 6C).

The diploidised auxiliary cell cuts off a first

gonimoblast cell which divides again (Fig.

6P) to initiate several branched gonimoblast

filaments with the ntature filaments terminating

in single carposporangia (Fig. 6£) which

mature simultaneously. New gonimolobes are

produced from the base of the gonimeblast and

mature Inter, Some darkly-staining and possibly

nutritive cells occur around the base of the old

auxiliary mother cell.

At an early stage in development of the pro»

carp. cells are cut off from the surrounding

cortical cells te form the projective pericarp

(Fig. 6D). The inner cells of the pericarp form

wz reliculum (the “tela aruchnonea™) which is

gradually absorbed by the developing carpo-

sporophyte (Fig. 6£), and the outer 2-3 layers

remain as the cystocarp wall, with a distinct

apical ostiole.

Spermatangia

Spermatangia are formed as in other species,

with small cells being cut off around the mar-

ging of the cortical ceils and then producing

branched chains of spermatangial mother cells

over the surface, froni whieh the clougate

spermatangia develop.

Tetrasporangia

The tetrasporangia develop by enlargement

of cortical cells (Fig. GF) which protrude

within the cortical layer, and they divide tetra-

hedrally (Fig. 6G).

Champia parvula (C. Agardh) Harvey 1853:

76. J. Agardh 1876; 303. De Toni 1900hb;

358, Newton 1931: 439, fig, 263. Gayral

1966: 485, pl. 134.

Chondria pervela C. Agardh 1824: 107,

Chylocladia parvala (C. Agardh) Hooker. Hur-

wey 1849; pl. 210,

Type loeality. Gades (Cadiz), Spain.

Type. Herb. Agardh, LD, 26(122,

Disizibution. C, parvula appears lo be the

only species of Chumpia known from Enropean

coasts, and hus been recarded from mast tem-

pefate and tropical coasts of the warld.

20 D. J. REEDMAN & H: B, S. WOMERSLEY

00 am

Fig. 6, Champia zostericola, A. Supporting cell with auxiliary cell branch and carpogonial branch

(443356). B, An early post-fertilisation. stage showing fusions between cells of carpogonial

branch (A43556). C. Post-fertilisation stage with fused carpogonial branch and connections to

auxiliary cell (A43556). D. Young gonimoblast within developing pericarp (A43556), E, Older

carposporophyte with terminal carposporangia, within pericarp (ostiole not in section)

(A43556). F. A young tetrasporangium (A34256). G. Mature tetrasporangium (A34256).

SQUTHERN AUSTRALIAN CHAMPIA AND CITYLOCLADIA $1

The following references credit C. parvaeda

to southern Australia, but probably all apply

to Other species, mainly (0 small specimens of

C. zostericola; ii Most cases it is Hot possible

to clarify these references

Guiler 1952; 94. Harvey 18552:

307. Lucas 1909: 34; 1929a: 19;

Lucas & Perrin 1947: 206, fig.

362. Reinbold 1897; 43. Souder” 1846: 176;

(S80; 17. Tate L832; 18, Tisdall ¢898; Sf,

Wilson 1892: 180.

C. parvula is generally recognised as a rela-

lively small and variahle species, and herbar-

ium specimens credited to it vary in size,

degree of branching, proportions of the seg-

ments and distinctness of the diaphragms, and

in the size of the cortical cells and degree of

outlet cortical development. Harvey (1849, pl-

210; (853, p. 76) commented on the variability

of this species, In comparisons with southern

Australian taxa, liquid preserved material from

He Verte, Rascoff, France (/. Feldmann,

14.2.1974; ADU, A46057) has been taken us

Tepresenmting the species in western Europe.

This specimen has fonger segments. (about as

long as broad) than shown by Gayral (1966,

pl, #34) bul distinctly shorter than illustrated

by Harvey (1849, pl. 210), and the dia-

phragms snd cortical cells are distinct through-

out the plant, with relatively slight development

of small outer cells. There are usually two com-

plete lonintuditial filament cells belween ihe

diaphragms, and the fongitudinal filaments are

confined to the periphery of the diaphragms.

The most detailed account of C. parvule is by

Bliding (1928) who studied material from

Woods Hole, ULS.A. Bliding’s deseriptiog and

illustrations appear to agree with the Le Verte

spectinen-.

In spite of the several references to C. par-

vale in southern Australia, it now seems clear

thal typical forms of this species do not occur

here. Young and small! plants of C. zoverfcole

do show some similarities, but are generally

broader, more robust, and when mature have

numerous hooked branches. Also. €. zosteri-

cola his usually only one complete longitudinal

filament cvll between the diaphragms, and the

cortical cells are lurger, Some small forms of

C. affinity also approach C. parvula but can be

Wistiguished on their greater outer cortical

development. Harvey's specimens referred to

C. pavvula appear to be slender forms of e:ther

C. zosterivola ar C, viridis.

However, a distinctive taxon occurs epi-

phytic on Amiphibolis at Tipaca reef in Spencer

Gulf, South Australia, and it appoars best to

S45, 1859;

1929b: SO,

70. May 1/9658:

designate this as a distinct variety of C. par-

vala, ta which it seents more closely allied than

to the larger €. gostericola. Fulure stuches may

show that tt should be recoynised as 4 distinct

species.

Champia parvula var. amphibolis vur. nov.

FIGS 7, 144

Thallus (Fig. 144) erect, spreading, 3-11

cm high, with one to several much-branched

Maiti axes arising from a small discoid hold-

fast on stems of Amphibolis, red to red brown

in colour, adhering to paper; oceasional attach-

ments by haustorial pads occur, Aves densely

and iegularly radially branched to 3 or 4

orders. with alternate, opposite or necamonally

whorled branches: axes 1-14 mm in diameter

below, tapering gradually to branchlets 13-4

mm in divmeter. Young branches slightly com-

stricted at «liaphragms, segments 1-14 times as

lung as broad, branch ends usually straight bud

rarely hooked, apices rounded. Dlapiiragens dis-

tinct throughout mosc of the thallus, somewhal

obscured near bases of older plants, Cortex of

a layer of angular cells 25-40(-50) pm across

and ¢1-)2-3(—4) times as long as broad, with

small cells cut off from their corners, and on

older axes developing a continuous guter cor.

lical Jayer (Fig, 7D) which in ol! plants may

be 3 cells thick. Longimdinal filaments con-

ffned to periphery of diaphragms, Ueveloped

from 10-15 apical cells, usually with evo (-3)

complete cells and two part cells between the

diaphragms (Fig. 7C).

Cystucarps single, subspherical to urceolate,

scaltered over younger branches, 27-4 mm

long and 4-1 mm in diameter, ostiolate.

Spermarengia Lorming son Over several seg.

ments near the apices of young branches,

Tetrayporangia scattered in young branches,

75-120 um in diameter,

Thallus erectus, effusus, 3-11 cm ialtus ex hap-

tere parvo discoideo in Amphibole. Axes irregu-

eulanter ramesi, 1-14 mm in diametro, ramuli

1/3-4 mm in diametro; segmenta I-14 plo

longlora quam fata, Diaphragmota conspicua nisi

prope bascs plantarum veterum, Cortex compositus

cellularum angulosarum 25—40(—350) #m Jalarum,

(1-)2-3(-4) plo longiorum quarm Jataram, parvas

cellulas in angulis ferens, ad 3 cellulas crassus in

parlibus veteribus crescens, Filamenta longitudin-

Alia tantum in margine, 2(-3) cellulas tolas inter

diaphraymata habentia. Cystocarpiy subgloboss vel

ureealjita, dispersa, ]_1} mm longa et + J mm itt

diametro, ostiolata, Spermatangla in soris fasci-

culatu prope apices ramulorum, Tetrasporansia

75-120 em in diametro dispersa.

Tig. 7,

D, J. REEDMAN & H. B. 8. WOMERSLEY

100 um

Champia parvula var. anphibolis, A. Surface view of branch showing apical cells (A41276).

B. Cross section of brunch showing diaphragm, cortex and peripheral longitudinal filaments

(A41276). C, Longitudinal section of a branch showing single-layered cortex, diaphragms, and

longitudinal filaments with gland ceils (A4J276). D. Longitudinal section of older axis show-

ing two (—3) layered coriex (A37291), E. Post-fertilisation stage showing fusions between

carpogonial branch cells and connection to auxiliary cell (A38255). F. Young gonimoblast

(A38255). G. Mature cystocarp {ostiole not in section) with carposporophyte bearing terminal

carposporangia (A38255). H. Section of male thallus with spermatangia (A41276). I. Mature

tetrasporangium (A37291).

SOUTHERN AUSTRALIAN CHAMPIA AND CHYLOCLADIA o3

Type locality, Tipara reef, Spencer Gulf, 5,

Aust. (Shepherd, 23,xi.1970)-

Type, ADU, A38255.

Distdibution. Knows from several collections

{rom Tipara reef, on Amphibelis antarctica

and A, griffithii, 5-11 ov deep. Young plants

occur ih June, maturing to bushy plants up to

11 cm high in December.

Var. amphibelis resembles the more typical

forms of Champia parvila in being a relatively

slender plant, mostly irregularly alternately

hanched, with diaphrapms distinct throughout

most of the plant ancl segments as long to

slightly longer than broad, with the primary

conical layer of cells cutting off relatively Few

outer cells (except in old parts}, and with

usually two complete longitudinal filament cells

between the diaphragms. Reproductively it ts

similar to the account of Bliding (1928) and

at least superticially to the Tle Verte material.

It differs from European forms in being slen-

derer than some, more profusely branched, and

in growing on Amphibolly as erect tufted

plants. Future calleclions may show that it is

not confined to this sea-prass and some speci-

mens of Harvey (in TCD & MEI.) from Fre-

mantle and King George Sound, W. Aust. (e.2.

Alz Aust. Exsice. 254B in MEL, 45307) need

careful comparison with this variety and with

C, zortericola,

This Australian variety shows slight simi-

larity to C. zestericola but is very much slen-

derer than most plants of the latter, only tarcly

has hooked branches, has longer segments close

10 the apices and has two complete longitudinal

filament cells between the diaphragms rather

than the usual one in C, zestericola. C. zosteri-

cola docs occur on Amphibolis, but is more

churacteristically associated with Postdenia.

STRUCTURE AND REPRODUCTION

Materia! studied: Tipara reef, Spencer Gulf, S.

Aust, on Amphibelis, Shepherd. The type, Ad1276

(13.xi1,197L) and AS72940 (30.14.1970),

Thallus development

The thallus has 10-15 apical cells (Pig. 7.4)

which segment us in other species to form a

peripheral ring of longitudinal filaments (Fig.

78). with diaphragms forming usually from

each third cell and thus with two (occasianally

three) complete Jongitudinal filament cells

{usually each with w gland cell} belween suc-

cessive diaphragms (Fig 7C). The cortex is

one cell itnck (Fig. 7C} except on older axes

(Fiz. 7D) and the elongate cortical cells cut

off, from their corners, small cells (nsually

only one per cell) which lic more er less in the

layer of primary cortical cells. These small cells

commonly bear a hair in young branches, but

such hairs are lost from older branches.

On old axes, the primary cortical cells cut

off an ovter, continuous, layer of cells (Fig,

7H) and this may become two or three Cells

thick, the outermost layer being of small cells,

Branching occurs from the regions of the

diaphragms, Mast branches are linear to che

apex, but occasionally a curved or slightly

“hooked” branch -end occurs.

The holdfast remains small and ciscoid, but

several new axes may develop from it, Small,

pad-like haustoria also develop from branches

of same plants, attuching mainly te other

branches.

Precarp and carpasporaphyte

The multinucleate supporting cell develops

from a cortical cell and hears hoth carpogonial

rnd auxiliary cell branches as in other species.

Following fertilisation, {he carpogonial branch

cells begin to fuse (Fig. 7E) and a connection

forms between the fertilised carpogonium and

the auxiliary cell. The diploidised auxiliary cell

forms 4 first ponimoblast cell which divides

again (Fig, 7E), and the upper cell forms the

branched gonimoblast filaments (Fig. 7G)

which bear terminal catposporangia.

Following fertilisation, some vegetative cells

arqund the supporting cell become darkly-

staining, apparently as nutritive cells, Other

Vegetative cells divide to form the pericarp

(Fic. 7G) which develops as in other species,

with the inner eclls forming the “tela arach-

noidea’ which is broken or absorbed by the

developing carposperephyte, A well-defined

ostiole occurs at maturity of the cystocarp,

Sperimatangia

The small ouler cortical cells, or Further cells

cut off from the primury cortical cella, divide

to form branched filaments of cells covering

the surface of one to several segments close to

the apices of young branchlets, Each cell of

these filaments functions as a spermatangial

mother cell which cuts off outwardly 2-3

clongate-ovold spermatangia (Fig. 7H), which

appear to be shed entire.

T etrasporangia

Tetrasporangia (Fig. 7/) develop within

cortical cells which enlarge greatly ane holge

within the segments of the thallus.

54 D. J. REEDMAN & H, B. S. WOMERSLEY

CHYLOCLADIA Greville

Chivlocladia Greville (in Hooker 1833, p,

297). with the type species C, kuliformuis, is

conserved over Kalifermis Stackhouse 1809,

and is distinguished from Cfiampia by the for-

mation of carposporangia directly from a large

basal tusion cell (without branched ponimo-

blast filaments as in Champia) and by the

eystocurps being non-ostiolate. Otherwise,

Chyloctadia is similar structurally to Champia.

While numerous Australian taxa have at

sume lime been referred to Ciivloctadia, most

have been placed in uther genera (see Kylin

1931) or can now (see below) be excluded

from Chvloclacia.

However, 3 very distinctive species of Chy/o-

eladia, known From only a few deep-water col-

lsctions, has recently been discovered in South

Austcalian walers.

Chyloctadia grandis sp, fov.

FIGS & S$, 14B

Thallus (Fiz. 148) crect, red-brown to red-

purple, 20-30-cm high, with one to several axes

urising from a hard, branched, perennial base

to 5 ¢m high and 1/3-¢ cm thick, attached to

rock by a discoid holdfast to 14 cm across,

Axes with opposite or usually whorled lateral

branches ta 10 cm Jong, similarly branched

{muinly oppositely or alternately) to a second

or third order; axes often denuded below, 2-7

mm in diameter, branches 14-24 mm in dia.

meter tapering to 2-1 mm in diameter in

branchlets; all branches slightly basally con-

séricted, tapering gently to a rounded apex,

segments 4-12 times as long as broad:

htanches slightly constricted between segments,

but diaphragms conspicuous. Cortex in branch-

Jets | cell thick (Fig. $0), in older branches

thicker and in axes to 8 cells. thick {Fig, S£):

cortical cells ovoid, 75-35(—40) zm across and

1-2 times as long as broad in surface view,

Longitudinal filaments Scattercd throughout

diaphragins, with (I-)2(-3) complete cells

und two part cells between the diaphragms

(Fig. 88), with each peripheral filament cell

connected to the cortical cells by a lateral fila-

merit.

Cystocerps scattered over lesser branches,

spherical to shghily ovoid, #2(-14) mm

across, broad based. without an ostiole; carpo-

sporangia borne directly on the large, basal,

Fusion ceil,

Spermatangia unknown.

Tetrayporangie scattered over branches,

ietrahedrally divided, mostly 150-200(—250)

pm ip diameter.

Thalluy erecins ad 20-5U cm alias unc vel

Pluribus axibus ex base dura rarnose e1 perconl ad

5 em altis et L/3-4 cm latis ortis, haptere dis-

coideo, Rami laterales in verticillum vel opposire

dispositi, ad 10 cm longi ct similita ramtosi: axes

2-7 mm, rami 1.5-2.5 mm et ramuli 0.7-£ mm in

diametro, segmentis $-14 plo longiorlbus quam

lalis, plus minus constrictus, diaphragmatibus con-

spicuis, Cortex ad nnam cellulam in ramulis

cyassus, ad 8 cellulas crassus in axibus crescens.

Filamenta fongitudinalia dispersa, plerumque 2

cellulas totas inter diuphragmata habentia. Cysto-

carpia subglobosa 3-1(=14) mim Inta, haud osteo-

lata, dispersa; carposporangia in walescenti cellula

ipsa magna basali portata. Tcirasporaneta in ramis

dispersa, 150-200(—250) em in diametro.

Type locality, Tapley Shoal, Edithburg, §,

Aust., 15 m deep (Shepherd, 2,11,1969),

Holotype. ADU, AS33515. Isotypes to be dis-

iributed under this number.

Distribution. Only known from the type col-

lection from Tapley Shoal, and Investigator

Strait, S. Aust., Wetsen, 11 m deep (WLIO I:

ADU, A40995}, 23 m deep (28.21.1971; ADU,

A41010), and 34 m deep (201-1971; ADU,

ASSL9T),

Chylocladia grandis appears to be quite dix

tinct in fs form, large size, detise branching,

and in the perennial base which appears 10 last

fov several years, producing one to several

fronds annually (probably in spring and Sasting

Uirough summer).

STRUCTURE AND REPRODUCTION

Material studied: The type and Investigator

Strait. collections.

Thallus development

The multiaxtal apex of a branch (Fig,

84, B) includes both a central group of apical

cells which give rise to the scattered longitud-

inal filaments, and outer apical oglls which pro-

duce the peripheral longitudinaf filaments and

the cortex.

Fig, 8. Chylocladia grandis, A, Surface view of an apex showing peripheral and central apical cells

(A33515). & Longitudinal section of a branch apex showing development of cortex, dia-

phraems, und longitudinal flaments with gland cells (A335145). C. Cross section of a branch

showing a diaphragm with sub-peripheral and central longitudinal flaments (A33515). D,

Longitndinal view of outer part of a mature branch, showing the lateral connecting filaments

between ihe longitudinal filaments and (he vortex

(AS35151. £, Lonintudinal section of an

axis showing the multi-layered cortex (A33515),

SOUTHERN AUSTRALIAN CHAMP/A AND CHYLOCLADIA

FIG. 8.

94 D, J. REEDMAN & H. B. & WOMERSLEY

acne Or

Fig. 9. Chyloeladia grandis. A, Longitudinal section showing supporting cell with auxiliary mother

cells und 4-celled carpogonial branch (A33515). B. Post-fertilisation Stage showing two auxiliar

cell branches and remnants of carpogonial branch (A33515). C. Post-fertilisation stage wit

auxiliary cells bearing young catposporangia, The ‘supporting cell, auxiliary mother cells and

auxiliary tells. are partly fused (A33515

), 2. A cystocarp with mature carposporangia formed

directly from the fusion cell, the lobes of which probably indicute the original auxiliary

mother cells (A33515). E. A young tetrasporangium (A39197 ). F. A mature tetrasporangium

(A39197).

The 12-16 central apical cells (Fig, 8A, B)

divide transversely and the cells elongate to

form longitudinal filaments. Each third ¢écll

usually produces a whorl of diaphragm initials

which divide further to join with similar

adjacent cells to form the single-layered dia-

phragms (Fig. 8B, C). The longitudinal fila-

ment cells between the diaphragms generally

produce a single spherical to slightly pyriform

gland cell (Fig. 8B).

The 15-20 outer apical cells divide trans-

versely to form the peripheral ting of Jongilu-

dinal filaments, but each of these cells, close ro

the branch apex, divides periclinally to form an

ouler primary cortical cell initial (Fig. &8).

This cell divides periclinally again once or twice

and then the outer cell divides anticlinally to

form the primary cartical layer, but the Later

formed cells are not in pit-connection with the

longitudinal filaments (Fig, 8B). The periclinal

division of the cortical initial is followed by the

inner one or two cells elongating to form a

bridging filament between each cell of the peri-

pheral longitudinal filaments and the cortical

cells (Fig. 8D), This feature is not found jn

the Australian species of Chantpia. The peri-

pheral longitudinal filament cells cut off dia-

phragm cells which joitt with those from the

inner filaments. The peripheral filaments are

usually separated by one diaphragm cell from

the cortex (Fig, 8C),

Many of the primary cortical cells cut off 1

small outer cell which produces a hair; these

hairs form a dense felt over most of the thallus.

SOUTHERN AUSTRALIAN CHAMPIA AND CHYLOCLADIA 97

pel eA VN a a a

1 2 3 4 > 6 7 8 9 10

Fig. 10. Champia viridis. A. Type of C. tasmanica Harvey in TCD. B. A slender form (var, gracilis

Harvey) on Posidonia (Marino, S. Aust. Drift. Womersley, 26.x.1975; ADU, A46651).

98 D. J. REEDMAN & H. B. S. WOMERSLEY

As the branch matures, the primary cortical

cells cut off outer layers of cells, and in old

axes the cortex may be up to 8 cells thick

(Fig. 8E).

Branches originate from the region of the

diaphragms, when a group of apical initials

develops from the cortical cells,

The basal part of the thallus is clearly peren-

nial, being hard and resistant, up to 5 cm high

with irregularly placed, lateral propections rep-

resenting the bases of previous axes. From the

number and position of branch bases, some

perennial bases appear to be 4—5 years old, and

the axes are probably newly formed in spring

and lost by the following winter. The branches

probably decay rapidly since this plant has

never been collected in the drift.

Procarp and carposporophyte

The supporting cell is a large primary cor-

tical cell in pit-connection with a peripheral

longitudinal filament (Fig. 94). The support-

ing cell is multinucleate and cuts off outwardly

a small cell, the carpogonial branch initial, and

two larger cells, the auxiliary mother cells. The

carpogonial branch (Fig. 9A) is 4-celled,

curved, with an outwardly directed trichogyne.

Prior to fertilisation, each auxiliary mother

cell produces a uninucleate auxiliary cell (Fig.

OB).

Following fertilisation, a connection forms

between the fertilised carpogonium and each

auxiliary cell, and carposporangia are formed

directly from the auxiliary cells (Fig. 9C).

Fusion occurs between the auxiliary cells and

auxiliary mother cells (Fig. 9D), forming a

large basal fusion cell bearing the carpo-

sporangia directly. The supporting cell and

some vegetative cells may be incorporated into

the fusion cell.

Concurrent with the early development of

the carposporophyte, vegetative cells around

the supporting cell divide to produce erect

chains of cells which cut off outer cells and

form the pericarp (Fig. 9D), as in Champia.

The inner cells of the pericarp form the “tela

arachnoidea”, but no ostiole is produced.

When the carposporangia are mature, the top

of the pericarp ruptures.

Tetrasporangia

The tetrasporangia develop by enlargement

of the primary cortical cells (Fig. 9E), which

have several pit-connections with adjacent cells.

The tetrahedrally divided sporangia (Fig. 9F)

develop a thick gelatinous sheath.

Relationships

Chylocladia grandis agrees well with

Chylocladia and its type species, C. kaliformis,

in thallus structure and in reproduction. How-

ever, it has not been established whether one

or two auxiliary cell branches occur in C.

kaliformis. Champia has only one auxiliary

cell branch as far as is known, but the type of

Gastroclonium Kuetzing (G. ovale (Hudson)

Kuetzing) has two auxiliary cell branches

(Bliding 1928, p. 27).

The thallus structure of Chylocladia grandis

differs from that of the Australian species of

Champia in that the initial cells of the primary

cortex are connected to the longitudinal fila-

ment cells via a filament of one or two cells,

not directly, Whether this occurs in C. kali-

formis has not been established,

SPECIES EXCLUDED FROM

CHYLOCLADIA

Apart from the various Chylocladia names

which have been shown previously to be syno-

nyms of species of Champia or other genera,

the following names are now referred to other

genera.

Chylocladia fruticulosa (Reinbold) De Toni

1900b: 576.

Lomentaria fruticulosa Reinbold 1899: 46,

Type locality, Investigator Strait, S. Aust.

(Davey 148).

Type. Herb. Reinbold, M. Isotype in ADU,

A1553.

The thallus of the isotype is on Posidonia

(not Amphibolis antarctica as in Reinbold). It

is hollow and without diaphragms, and the

tetrasporangia are grouped in sori around

depressions in the wall of the branches, These

features are typical of Lomentaria, and the iso-

type (a small, bleached specimen) appears

similar to the earlier described Lomentaria aus-

tralis (Kuetzing) Levring 1946, p. 223 (Chon-

drothamnion australe Kuetzing 1865, p. 29, pl.

82 d-f). The southern Australian species of

Lomentaria are in need of detailed study.

Chylocladia gelidioides Harvey 1863, synop.:

46. De Toni 1900b: 578; 1924: 312. Gepp

& Gepp 1906: 257. Okamura 1904: 88.

Type locality. Twofold Bay, N.S.W. (F.v.

Mueller).

Type. Herb. Harvey, TCD.

Although cystocarpic material has not been

studied, the hollow thallus construction without

single layered diaphragms, and sori of tetra-

sporangia, are typical of Lomentaria. The

z t

HT!

€

& 8 Z 9

Hecrvabiinfrcalicatcand sotirnlcatitanitosttuitnaoadatativt

SOUTHERN AUSTRALIAN CHAMPIA AND CHYLOCLADIA

22-6.

a WA ‘in iq OTL CL WL Uy re

2 8 ! | ! ei ;

it 1

Fig. 11. A. Champia insignis. Isotype male specimen (ADU, A12237).

B. Champia affinis. Lectotype specimen in BM.

6 7.

oe he eae

100 D. J. REEDMAN & H. B. S. WOMERSLEY

[rset wa daa ae WL

Thhal

Re a 2 A

sa, hey a coe Lad

1 2 3 4 5

> - ‘

“a f or —

i rete ye fa Ft ttn eets oe tes

Fig. 12. Champia affinis. A. Harvey’s Alg. Aust. Exsicc. 2521 (as C, obsoleta) from Georgetown, Tas.

(in TCD). B. Lectotype of C. obsoleta (Port Fairy, Vic. Harvey’s, Alg. Aust. Exsicc, 252D)

—a rough-water form. Champia zostericola. C. Harvey’s Alg. Aust. Exsicc. 249A, Fremantle,

W. Aust.

SOUTHERN AUSTRALIAN CHAMPIA AND CHYLOCLADIA 101

‘ P /

A |

ae 4 | ' ,™

tte TE 4 4 aed

oe | \ ad ad ; aj

? . Fy bee Lis

: & '

1 i : Ss ‘ Ps

. ‘ yy -, Ms, i e =— 2

q. TZ AN j ~ a =

a. ap eo te te * ah ew, Ae 2

Phe (je ot ade, OO: Loe a Te ae =-o

LPass, 3” a | au - aa 4 A? oo sf =

Sy F % 4 =

hE, ORR Lee ah

it~ ® : ae fr =a os t f sf =

(3S ewe 5 i ee ~ =

‘ * id —

=—wn

=—

—

8 é 9 s 7 e€ €

tow svt gtiuatuadustuniduatustihuslt

Fig. 13. A. Lectotype of C. affinis var. arcuata Hooker & Harvey. Gunn 1332, in BM. B. Champia

zostericola. Musselroe Bay, Tas. Perrin, March 1937 (MEL 45252)—plant with well-

developed axes.

102 D. J. REEDMAN & H. B. S. WOMERSLEY

"?,

AON

a eee ww Ie

Be 7h

ie aan MM

1 2 3 4 5 6 7 8

§ A. Shepherd

toll

Fig. 14. A. Champia parvula var. amphibolis. Type specimen of variety.

B. Chylocladia grandis. Holotype specimen.

SOUTHERN AUSTRALIAN CHAMPIA AND CHYLOCLADIA 1%

thallus habit, cell detail, and tetrasporangial

sori are very similar to Lomentaria catenara

Harvey from Japan (as noted by Harvey 1863

and Okamara 1904), and the N.S.W. plant

may be a slightly less robust formi of the

Japanese species, Which is ulso recorded from

Pacific Mexico by Dawson (1963, p. 465, pl-

92). It appears to by closely related to Lomen-

faria ramsayana (J. Agardh} Kylin (1931, p.

27, pl, 14, fig. 33),

Chylocladia multiramea Sonder 1853; 681,

Type locajity, Lefevre Pen., S. Aust-

Type. MEL, 45196,

The type specimen (female) in MEL is a

slender, much branched, bleached plant of

Dasyphiloea insignis Montagne.

Acknowledgements

The authors are indebted to the Directors af

the National Herbaria of Victoria and of New

South Wales, and to the Brifish Museuns

(N.H), London, for the loan of specimens,

Professor J. Feldman, Institut de Biologie

Veégetale, Paris, Kindly seot matenul of

Champia parvula from Brest, and Mr R. H.

Simons, South African Seaweed Research

Laboratory, forwarded material of C. lnmbri-

caliv and C, conpressa,

The second author gratefully acknowledves o

grant from the Australian Research Grants

Committee and the technical assisiance pro-

vided by Mrs Enid Robertson and Miss Chery!

Anderson.

References

Acarpy, C. A. (7824),—“Systema Algarum,”

(Lund.)

AGAroH, C, A. (1828) —'Species Alvarum." Vol.

2. (Grypbiswald.)

Agarpy, J. G. (1852),—"Species, Genera ct

Ordines Algarum.” Vol. 2, Pt 2, pp, 337-720,

P\ 3, pp, 7O1-7E6. (T.und.)

Acaron, J. G. (1876).—"Species, Genera et

Ordines Algarum.” Vol, 3, Pri, pp. 1-724—

Epicrisis systematis Floridearum. (Lund,)}

AGABDH, J. G, (1879).—Florideernes morphologi,

K. VPerensk Ake. Forsehanedl, 15, 1-199,

Plates 1-33.

Acagoy, J, G. (1901).—"“Species, Genera et

Ordines Algarum, Vol, 3, Pr 4, pp. 1-149.

{Lund.)

Biome, C. (1928),—Studien tiber die Florideen-

ordoung Rhodymeniales, cfa Univ. aud,

N.F. Avd, 2, 24(3), 1-73.

Cuapman, V. J,, & Dromaoo.e, FT. (1970) -—

The marine alese of New Zealand. Pa7t UT:

Rhodophyceas. Florideophycidae; Rhodymme-

hiales. pp. 185-(54, Plates 39-50,

Dawson, E, Y. {1963).—Marine red algae from

Pacific Mexico. Part 6. Rhodymeniales, Nova

Hedwigia 5, 437-476, Plates 77-95.

Desvaux, N. A, (1808),—Champia Desv, J, de

Bor, 1, 245-246.

De Toni, G. B, (1900a)—Tl genere Champia

Desv. Mem. Ponnf. Accad, Nuovi Lincei,

Roma 17, 65-80.

De Tont, G. 8. (1900b).—"Sylloge Algarum om-

nium hucusque Cognitarium.” Vol. 4. Floti-

deae. Sect. 2, pp, 387-776. (Padua.}

De Tont, G, B, (1924).—"Sylloke Algarum om-

nNipm hucusque Cognitarium.” Vol. 6. Plori-

deae, (Padua.}

Garnet, J, R. (1971) —Check-list No. fh—Marine

and Freshwater Algae. In “The Wildflowers of

Wilson's Promontory National Fark", pp, 93-

97. (Lothian: Melbourne.)

Gaynas, P. (1966) —“Les Aleues des Cétes Fran-

cases.” (Deren: Paris.)

Gepp, A. & Gere, BE. S. (1906}—Some marine

algae from New Sonth Wales, J, Bor, Lari,

1906, 249-261. Plate 481.

Gumex, E, R, (19523.—The marine alpue of Tas-

matua: check hist with localities. Pap, Prac.

R. Soc, Fasm, 86, 71-106,

Harvey, W. H. (1$38}.—"The genera of South

African plants,” (Cape Town}

Hanver, W. H. (18444).—Champia tasmanica; 2

new Australian alga, Lond. J, Ber. 3, 407,

Plate 19,

HakVey, W. H. (1844b).—Algse of Tasmania.

Land, J, Bor, 3, 428-454,

Harvey, W, H, (J847).—"Nereis Australis,”

(Reeve: London.)

Harvey, W. H. (1849),—"Phycologia Britannica."

Vol, 2, Plates 121-240. (Reeve; London.)

Harvey, W. H. (1853).—"Nereis Boreali-Ameri-

cana,” Part T],—Rhodospermae: (Smithsonian

Tnst.:. Washington.)

Harvey, W, H, (1855a),—Some account of the

Marine botany of the colony of Western Aus-

tralia. Trans. R. Irish Acad. 22, 525-566,

Harvey, W, H. (18555),—Algae. da J. D, Hooker,

“The Botany of the Antarctic Voyage", Part

lL. Flora Novae-Zelandiae. Vol. 2. pp. 211-

266, Plates 107-121.

Harvey, W. H. (1859),— Algae, dn J, D. Hooker,

“The Botany of the Antarctic Voyage", Part

TIL. Flora. Tasmaniae. Vol. 2, pp. 282-343,

Plates 185-196.

Hanvey, W. H. (1863).—"Phycologia Australica,”

Vol. 5, Plates 241-300, synop. pp. 1-73.

Hooxer, J. D,, & Harvey, W. HB. (1847) —Aleac

Tasmanicae. Land. J. Bor. 6, 397-417.

Heoneh, W. J, (1833)—"The British Flora,”

Vol. 2.

Jorr, A. B, (1965}—Plora Marinhe do litoral

norte do estado de Sao Paulo e regioes cir-

cunvizinhas, Bol, Foc, Files, Ciéac. Letras,

Univ. Sdo Paula, 294, Ser. Bal. 21, 7-395.

Kinc, R. I, Brace, J, Hope, & Durken, Sophie

(1971) —Port Phillip Bay Survey 2. 8. Inter-

tidal ecology of Port Phillip Bay with system-

wile Usis of plants and animals, Meyn, nein.

Mus. Viet, 32, 93-128,

Kuerzine, F. T. (1843)~-"Phycologia generalis.”

(Leipzig,)

Kuerzinc, F. T-.

( Leipzig.) .

Kuerzinc, F, T. (1865).—Tabnlae Phycologi-

cae.” Vol. 15. (Nordhausen.)

Kyiex, H. (1931) —Die Florideenordnung Rhody-

meniales, Acta Univ, lund, N.F. Avid. 2, 27

(11), 1-48, Plates 1-20,

Kyarn, H, (1956),—“Die Gattungen der Rhodo-

phycoen.” (Lund.)

Luvaing, T. (1946),—A list of marine algac from

Australia und Tasmania. Acta Herti gotho-

burg 16, 213-227.

Linnazus, C. (1771).—“Mantissa plantarum

altera.” pp. i-vi, 143-588. (Stockholm.)

Lucas, A. H. S. (1909).—Revised list of the

Fucoideae and Florideae of Australia. Proc.

Linn, Joe, N.S.W, 34, 9-60,

Lucas, A. H. S, (19294).—The marine algae of

Tasmania. Pap, Proc. R. Soc. Tasm, 1928,

f-27-

Lucas, A. HLS, (1929b),—A census of the marine

algae of South Australia. Trans. R. Soc: S$.

Aust, 53, 45-55. ;

Lucas, A. H. S. (1931)—Notes of Australian

marine algae, WI. Proc, Lina. Soc, NSW.

56, 407-411, Plates 23-27,

Lucas, A. H.S., & Pesnin, F. (1947).—"The Sea-

weeds of South Australis.’ Part TW. The Rexl

shih teal pp. 109-458. (Govt Printer: Ade-

aide,

May, V. (1947).—Studies on Australian marine

ulgue. IMT. Prac. Tian. Soc. N.S.W. T1, 273-

277, Plate 19,

May, V. (f965).—A census and key to the spe-

cles of Rhodophyceae (red algae) recorded

from Australia. Contr. N.S.W. narn. Herh. 3,

349-429

Navanir, C. (1847).—"Die neueren Algenstyeme,”

(Zurich)

Nayior, M. (1954).—A check list of the marine

algae of the Dunedin district. Trans, R. Sec,

N.Z. 82(3), 645-663.

Nrwrton, L, (1931),—A Handbook of the British

Seaweeds.” (British Museum: London,)

OKaAmuRA, K, (1904),—List of marine algae col-

lected in Caroline Islunds and Australia. Bot,

Mag., Tokyo 18, 77-96,

REINBOLD, T. (1897) —Die algen der Lacepede

und Gilichen Bay. Nueva Notarisia 8, 41-62.

KEINBOLD, T, (1898) —Die Algen der Lacepede

und Guichen Bay (Stid Australien) und deren

naherer Umbebung, gesammelt von Dr <A,

epgeihact-SoaiRsog: I. Nuova Notarisia 9,

33-54 .

(1849) —"Species Algarum,”

TD. J, REEDMAN & H, B.S, WOMERSLEY

REINBOLD, T. (1899),—Meeresalgen von Investi-

gator Street (Sud Australien), gesammelt von

Miss Nellie Davey (Waltham, Honiton). Hei-

wigia 38, 39-51,

SuEPHERD, S. A., & WoMERSLEY, H. B, S, (1970).

—The sublittoral ecology of West Island,

South Australia. I, Environmental features

and the algal ecology. Trans. R. Soc, S. Avast,

94, 105-138.

SHEPHERD, S, A., & Womerscey, H, B. §, (1971),

—Pearson Island Expedition 1969.—7. The

sub-tidal ecology of benthic algae. Trans. R,

Soc. 8S. Aust. 95(3), 155-167.

SonpDER, O. G. (1846).—Algae. Jn C. Lehmann,

“Plantae Preissianae,”’ Vol, 2, pp. 148-195.

(Hambureg.)

Sonper, O. G, (1853),—Plantae Muellerianac,

Algae. Linnaeu 25, 657-709.

Sonver, O. G_ (1855).— Algae annis 1852 et 1855

collectae. Linnaea 26, 506-528.

Sonper, O, G. (1880) —/w F, von Mueller, “Frag-

menta Phylographiae Austratiae". Supple-

mentum ad volumen undecinum; Algae Aus-

tralianae huctenus cognitae. pp, 1-42, 105-107,

StackHouss, J, (1809).—Tentamen Marino-

Cryptogamicum. Mem. Soc. Imp. Nat. Mes-

cou 2, 5097.

Tare, R. (1882).—A list of the charas, mosses,

liverworts, lichens, fungs, and algals of extras

tropical South Australia, Trans, R, Sec. §.

Angst. 4, 5-24.

Tispait, H. T. (1898).—The algac of Victoria,

Rep, 7th Meet. Aust. Ass. Ady. Sci., Sydney,

1898. pp, 493-516.

TREVISAN, B. A. (1848),—“Saggia di uns

monugrafia delle alghe cocotalle.” (Padova,)

WEBER VAN Bossg, A. (1928). —Liste des algucy

du Siboga. IY. Rhodophyceae, Pt IT, Gigar-

tinales «ct Rhodymeniales. Monogr. Siboga

Exped, 59d, 393-533, Plates 11-16.

Wirson, J. B, (1892) —Catalogue of algae col-

lected at or near Port Phillip Heads and

Western Port. Proc, K, Soc, Vier. 4, 157-190).

WoMERSLEY, H. B.S, (1948).—The marine algae

of Kangaroo Island. If. The Pennington Ray

region. Trans. R, Soc, 3. Aust, 72, 143-166,

Plates 10-15.

Womerscey, H. B, S, (1950).—The marine algae

of Kangaroo Island. If. List of species 1.

Trans. R. Soc. §. Aust. 73, 137-197,

Womersiey, H. B. S. (1966)-—Port Phillip sur-

vey, 1957-1963: Algae. Mem. notin. Mus.,

Vier, No, 27, 133-156.

WoMERSLEY, H. B. S., & Batty, A. (1970}.—

Marine algue of the Solomon Islands, PAii.

Trans, R, Soe. Lond, B 259, 257-352,

LEGGADINA LAKEDOWNENSIS, A NEW SPECIES OF MURID RODENT

FROM NORTH QUEENSLAND

BY C. H. S. WATTS*

Summary

WATTS, C. H. S. (1976).-Leggadina lakedownensis, a new species of Murid rodent from north

Queensland. Trans. R. Soc. S. Aust. 100(2), 105-108, 31 May, 1976.

Leggadina lakedownensis, a new species of murid rodent from northeastern Queensland is

described and figured. Morphologically it most closely resembles L. forresti (Thomas).

LEGGADINA LAKEDOWNENSIS, A NEW SPECIES OF MURID RODENT FROM

NORTH QUEENSLAND

by C.H. S, WaATTs*

Summary

Warts, C. H. S. (1976) —Leggadina lakedownensis, a new species of Murid rodent from north

Queensland. Trans, R. Soc. §. Aust, 100(2), 105-108, 31 May, 1976.

Leggadina lakedownensis, a new species of murid rodent from northeastern Queensland is

described and figured. Morphologically it most closely resembles L. forrest? (Thomas).

Introduction

A pair of small rodents from Cape York,

examined during the course of a broader inves-

tigation into the karyotypes of Australian

rodents, was found to differ in karyotype,

blood proteins, and cranial morphology from

the otherwise similar Leggadina forresti, as Well

as all other murids examined,

Leggadina lakedownensis n.sp.

FIGS 1, 2, and 3

Holotype: Qld Mus, JM1192, 9. Received

from Queensland Museum in May 1975, killed

August 1975. The parents of this specimen

were received by the Queensland Museum from

Mr C. Tanner in August 1973; these were off-

spring of animals collected at Lakeland Downs,

110 km S of Cocktown, Queensland by Mr R.

Buckley in 1973.

Description: (Colour after Ridgway 1912).

Head relatively narrow and pointed as in L.

deliculata and L. hermannshurgensis. Eyes. not

as prominent as in those species or in L. for-

resti. Ears small and broad, proportionately

smaller than in L. forresti. Feet narrow. Hairs

on back with tips buffy brown grading to pale

olive-buff on sides, bases of hairs blackish

brown which shows through giving back a

brindled look, Underside white, hairs white to

base, as are hairs on upper surface of feet, Face

with suggestion of darker central stripe and

lighter ring around eyes, Tail sparsely haired

with light-coloured hairs.

Skull flat on top, rostrum short, interorbital

region broad. Interparietal wide, short. Zyge-

Fig, 1. Dorsal, lateral and ventral views of skull of Holotype Leggudina lakedawnensis n,sp., % 2.5.

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

16 C oH, 3S. WATTS

Fig, 2. Upper right molar tooth row of holotype o}

Leggadina lakedownensis tsp,

matic plates moderate, minimum width equal

to Jength of Mt. Lacrimals small. Nasals short,

not exceeding premaxillae anteriorly. Incisive

foraminae longer than tooth row, reaching

heyond anterior end of M4, broad, slightly

Wider posteriorly, braadiy rounded posteriorly,

more pointed anteriorly. Posterior palatal fora-

minae oval, about lencth of M5. Mesopterygoid

fossae narrow, their width about equal to length

of Mi. Bullae moderately developed about size

of occipital foraminae, the distance between

them about equal to combined lengths af M=

and M3. Upper incisors forward pointing.

Molars as in Figure 2, anterior ligual cusp of

M} bDlade-like and strongly developed. M+ large,

approximately 659% of tooth row, M% small,

approximately 15%. of tooth row.

Specimens examined; Paratypes; Qld Mus.

JM1293 3. JM1294 f bred in captivity; collec-

tion details as for Holotype, Referred specimen,

Qld Mus. J.17919, Williams Id, Queensland

(35°S; 135°B), 1969, coll. Mr, C. Tanner

Diagnosis

(Head and body 60-70 mm, Tail 40-50 mm,

Far 10-12. mm, Skull length 20-23. mm),

Incisive foraminae reaching heyond anterior

end of M}, rounded and slightly widened pos-

teriorly. Upper incisors forward pointing.

Upper molar tooth row 3.7—4.2 mm long, An-

terior ligual cusp of M} blade-like and strongly

developed. M} 4 to 5 times length of M}.

Systematic position

Leggadina lakedownensis ts separated from

all named forms within the genus Legvadina, as

recognised by Tate (1951), except L. forrest

(Thomas), L. waite’ (Troughton) und LZ, mes-

soria {Thomas), by the short tail, small cars,

Jong incisive foraminac, and [arge molar tooth

row with M1 large and with a strongly

developed antenor ligual cusp.

These three taxa, L. forresti, L. waitei and

L. miessoria, were considered to he very closely

related by Tate (1951) and were synonymised

under the name of Pseudomys forresti by Ride

(1970). My examination of the types of all

three supports this view that only one species

is involyed, characterized by downward point-

ing upper incisors and long incisive foraminae

harrowing posteriorly and about 3 times the

length of M3. Measurements of the Holotypes

are included in Table 2-

Lepvadina lakedownensis differs from L. for-

resti (and the Holotypes of L. waifei and L.

messoriag) in having the incisive foraminae

widening rather than harrowing posteriorly, a

slightly larger M} (in comparison to total tooth

tow), a smaller M3 und forward pointing upper

invisors, On the limited evidence available it is

TABLE 1

Measurements o] specimens of L. lakedownensts (min)

1 2 3 4

Hend and body 72 - — 64

Tail 45 — -— 4)

Hind toot (S.u.) 1B _— _ MW

Far (from noteh) It a — i

Greatest length of skull 73.0 2s — aA

Zyyomatic breadth 125 tla 12.0 10.8

tnterorbital breadth a4 45 44 tn

Depth of brain case,

inclyding bullae R4 - _— 36

Length of nasal 73 73 73 6.7

Length of ant. palatal

foramina a7 5.2 54 4.8

Crown length of molar vow 4.0 49 3.7 4.2

Crown length of M1, 27 2.7 24 246

Crown length of MA 04 O58 0.5 09

1, Old Mus. IM1292 9. Lakeland Downs, Qld, 1973 Holo-

type.

2. Qld Mus. JM1294 9, Lakeland Dowhs. Ql, 1973

3. Old Mus. M1293 9, Lukeland Downs, Qtd, 1973,

4, Qld Mus, 317919, Williams Td, Qld, 1969,

NEW SPECIES OF MURID RODENT

107

Fig. 3. Leggadina lakedownensis, tsp.

a slightly smaller animal. In my opinion the

scale of these differences indicate a form speci-

fically distinct from L. forresti or any other

described rodent,

Support for the distinctiveness of L. lake-

downensis from L, forresti comes from studies

of the chromosomes and blood proteins of these

and some related species (Baverstock ef al.

1976). These studies suggest that L. forresti

and L. lakedownensis form a distinct group

within Pseudomys sensu lato. The morphologi-

cal characters (cf. above) tend to support this

view and it seems prudent at this stage to retain

the genus Leggadinu at least for these two

species.

Habitat and distribution

The Williams Island locality is an area of

short grassland surrounded by scrub. (The

locality is illustrated as Lakefield Station, by

Covacevich (1974, p. 7).) Lakeland Downs is

on an isolated area of basalt-derived red and

brown soil supporting a natural vegetation of

Box woodland (E£. leptophleba), and deciduous

scrub with kangaroo (Themeda australis) and

spear (Heteropogon contortus) grass. At the

time of collection however, sorghum covered

the whole area. Rainfall is between 100 cm and

130 cm per year, occurring mainly in the sum-

mer (J. Covacevich, personal communication).

At present the species is only known from

the above 2 localities on the eastern side of

Cape York. Leggadina forresri has a much

greater distribution in inland Australia and is

known from W.A., S.A., N.T, and N.S.W. as

well as western Queensland. From specimens

in the Australian Museum and Queensland

Museum (Table 2) it appears that there is a

considerable gap between the distribution of

the two species.

Acknowledgments

Miss Jeanette Covacevich kindly sent the

animals on which the descriptions are based.

Curators in the Queensland Museum, Austra-

lian Museum and South Australian Museum

kindly sent material or allowed me access to

specimens of Leggadina in their care. I am

also grateful to Miss Heather Aslin for draw-

ing the skull and teeth of the type specimen,

108

C. H. S. WATTS

TABLE 2

Measurements of specimens of L. forresti from Qld, S.A, and N.T. including Holotypes of L. waitei, L. forresti and L-

Head and body

Tail

Hind foot (S.u.)

Bar (from notch)

Greatest length of skull

Zygomatic breadth

Interorbital breadth

Depth of brain case,

including bullae

Length of nasal

Length of ant. palatal

foramina

Crown length of

molar row

Crown length of ML

Crown length of Mi

Baversrock, P. R., Hocartu, J. T., Core, S., & Ripeway, R.

(1976)—Biochemical and

Karyotypic evidence for the specific status of

the rodent Leggadina lakedownensis Watts.

CovacevicH, J.

AM M5194, Hart Range, N.T.

L. waitei Holotype

MNH 6.3,9.39, Alexandria, N.T., 1905

. forresti Holotype

a BMNH 25,4.9.1, Melrose, S.A,, 1922

— L. messoria Holotype

Trans. R. Soc. 8. Aust, 100(2),

Ripe, W. D. L. (1970).—"A guide to the native

mammals of Australia.” (Oxford Univ. Press:

London.)

x & AM M9208, Mt Isa, Qld, 1968

messoria (mm)

2 af ao

: - 5

o = Um =

; o

5° 27 s

3 3 2

e @ ¥g vg

3S cel mo ee

e 5 se 3&

= 5 ZG Zé

= = qt G<

< SF KH “nH

83 88 100

—_ — 5 6

172,-— 17 17

12.3 — 15 14

— 23.5 23.4 24.6

13.8 126 12.1 13.2

40 3.7 36 33

86 87 86 9.2

1.700 81

50 52 454 56

45 46 46 43

2.7 #27 27 25

09 09 10 O08

References

SAM M2958, Macdonald Downs,

S.A,, 1930

SAM M2405B, Horn Exp. (Waite)

SAM M2405, Horn Exp. (Waite)

SAM M6350, 60 km N Birdsville,

Qld, 1966

SAM M6351, 60 km N Birdsville,

Qld, 1966

SAM M6352, 48 km N Oodnadata,

S.A., 1966

(1912).—"Colour standards

colour nomenclature.” (Ridgway: Washing-

ton).

Tate, G. H. H. (1951).—Results of the Archbold

Expeditions. No. 65. The rodents of Australia

AM M9481, 20 km W Innamincka,

. 1973

and

and New Guinea. Bull. Am. Mus. Nat. Hist.

97, 183-430.

CovacevicH, J., & Easton, A. (1974).—“Rats and

mice in Queensland.” (Queensland Museum

Booklet No, 9: Brisbane).

BIOCHEMICAL AND KARYOTYPIC EVIDENCE FOR THE SPECIFIC

STATUS OF THE RODENT LEGGADINA LAKEDOWNENSIS WATTS

BY P. R. BAVERSTOCK*, J. T. HOGARTH*, S. COLE* AND J. COVACEVICHT

Summary

BAVERSTOCK, P. R., HOGARTH, J. T., COLE, S., & COVACEVICH, J. (1976).-Biochemical

and karyotypic evidence for the specific status of the rodent Leggadina lakedownensis Watts.

Trans. R. Soc. S. Aust. 100(2), 109-1 12, 3 1 May, 1976.

Leggadina lakedownensis Watts differs karyotypically from its apparent nearest relative, L. forresti.

Further, the biochemical differences between L. lakedownensis and L. forresti are greater than those

between other "good" species of similar sized pseudomyins. These data support the specific status

of L. lakedownensis.

BIOCHEMICAL AND KARYOTYPIC EVIDENCE FOR THE SPECIFIC STATUS OF

THE RODENT LEGGADINA LAKEDOWNENSIS WATTS

by P. R, Baverstock*, J. T. HoGARTH*, 8, CoLe* and J. CovAcEvicHt

Summary

Baverstoce, P. R. Hocarru, J. T., Core, §, & Covacevicn, J, (1976),—Biochemical and

karyotypic evidence for the specific status of the rodent Levvadina lakedownensis Watts.

Trans, R, Soc. §, Aust, 100(2), 109-112. 31 May, 1976,

Lexeadina lakedownensis Watts differs karyotypically from its apparent nearest relative.

L, forresti. Further, the biochemical differences between L. lakedownensis and L. farresti are

greater than those between other “good” species of similar sized pseudomyins, These data

support the specific status of L. lakedownensis,

Introduction

In 1973 several specimens of a species of

small rodent were collected from Lakeland

Downs in northeast Queensland and described

as a new species, L. lakedewnensis, by Watts

(1976), The morphological differences between

L. lakedownensis and its apparent nearest rela-

tive L. forresri are, however, minor. Because

speciation is the result of the accumulation of

many genetic differences and because mar-

phology alone reflects only part of these genetic

differences, it seems desirable in such cases to

assess. other aspects of genetic differences

between allopatric populations. The present

study was undertaken to determine whether the

karyotypic and biochemical differences between

L, forrestt and L. lakedownensis substantiated

the recognition of the latter as a distinct species,

Methods

Seurces of attimals: The sources of animals

utilized in the present study are shown in Table

1, along with their Institute of Medical and

Veterinary Science number, When these ani-

mals die their skull and skin will be submitted

to a Museum and given a Museum number.

Muscum numbers corresponding to JMVS

numbers. will be available from the IMVS or

the South Australian Museum,

Chromosome preparations; Chromosome

preparations were made from & L. forresti

and 3 L, lakedownensis. Animals were bled

by cardiac puncture under ether anaesthesia

and leukocytes cultured for 3 or 4 days.

Slides were prepared by means of the routine

TABLE i

Rejerence numbers and sources of animals

IMVS

Species No. Sex Locality

(a) Chromosomes

L. jarresti 1 EF Coorabulka, SW Qld

2 M = 19km W Innamincka,

S\A.

3. FEF Wk W Innamincka,

S.A.

5 F 19km W Innamincka,

SLA.

6 M ~ Bowlers Gop Stn, N.S.W.

7 \M ~~ Fowlers Gup Stn, N.S.W

8 M_ Fowlers Gap Sin, N.S.W,

9 M = Mt Sarah Sta, S.A.

L. lakedownensis 10% F Lakeland Downs Stn, Glu

11 Ms Lakeland Downs Stn, Qld

12 M_~ Lakeland Downs Stn, Old

(b) Blectrophoresis

L. forresti 3 fF 19km W Innamincka,

§.A.

5 FB 19km W Innamincka,

5A.

L, lakedawnensis 10* F Lakeland Downs Stn, Qld

L. delicatule 13. M_ Fairbairn Dam, 22km

SW Emerald, Qld

14 M_ Fairbaim Dam, 22 km

SW Emerald, Qld

L, hermanns-

burgensis 14 M_~ Lah, stock

P.novaehollandiae 15 M Port Stevens area, N-S-W,

P. australis M sLab. stack

* Holotype—Queensland Museum JM{292-

* Institute of Medical & Veterinary Science, Frome Rd, Adelaide, S. Aust. S000.

| Queensland Museum, Gregory Tce, Fortitude Valley, Qld 4006.

110

P. R. BAVERSTOCK, J. 1, HOGARTH, S, COLE AND J. COVACEVICH

TABLE 2

Eléctraphoreétic hiiffer atid Staitiing systeiis used for proteins examined

Enzyme/ Protein

E.K. t.1.149 Glucose 6-Phusphate dehydrogenase (G 6-PD)

6-Phosphoglicondle dehydrogenase E.C. 1.) 1.44 (6-PGD)

Phospho-hexose isomerase F.C. 5.3.1.9, (PHI)

Phosphoglucomutase E.C, 2;7.4.1, (PGM)

Lactate dehydrogenase E,C_ 1.1,1,27 (LDH)

NAD-Malhiute dehydrogenase E.C. 1.11.37 (NAD-MDH)

Leucine aminopeptidase E.C, 3.4.1.1 (LAP)

Glyceraldchyde 3-Phosphate dehydingenase GA 3-PD EC, 1,2.1.12

Telrazolium oxidase

Esterase (Est)

Albumin (Alb.)

Heccmich Asie { Ris;

Transferrin (TF)

* PEB — Tris-EDTA-Borate

air-dry method, For karyotypes, slides were

Stamed with 2% Giemsa, C-staining was con-

ducted by the method of Arrighi and Hsu

(1971) except that the RNase step was omitted.

Slides were then stained with 10% Giemsa.

Electrophoresis: In addition to 4. forrest! and

L, lakedownensis, specimens of L.. delicatula, L.

hermansburgensis, Pséudomys noyachollandiae

amd P. australis were studied, Blood was col-

lecred by cardiac puncture umder ether anaes-

thesia in syringes containing a dried film of

heparin and centrifuged immediately at 2,.000G

for 10 minutes at room temperature. Plasma

was pipetted off and stored at ~ 20°C. The red

cells were washed 3 times in 2 volumes of

isotonic saline and lysed in ai equal volume

of distilled water containing [/5 volume of

toluene. Cell walls were centrifuged out and

the clear supernatant stored at —20°C for a

Maximum period of 3 weeks.

Horizontal starch gell electrophoresis was

used. Gel slabs 300 mm x 150 mm x 6 mm

were prepared from 12.5% (50g/400 ml

buffer) starch using a perspex mould. Gels. were

run inva refrigerator to minimize heating. After

electrophoresis a section approximately 90 mm

x 120 mm was cut from the gel slab, sliced into

twa separate halves, and these halves then

Stain

Brewer (1970)

Brewer (L97/4))

Brewer (1970)

Buffer system

Brewer (1970)

Selander e7 ai, (1971) 7

Selander ef al. (1971) 7

Holmes ef a/, (1973)

TEB*

Holmes etal. (1973) Brower (L970)

TER"

Selander ef af, (1971) 2 Rrewer (1970)

Brewer (1970) Brewer (1970)

Brewer (1970) Scored from gels

stained fram GA

3-PD

Soin A; 008M. tris-cit- Brower (1970)

rate pH 8.6 Soln Br

0.06 M. Li-borate-pH

3.3 Gel: 337.5 ral Soln

Ar 62.5 ml Sola B in

400 ml B. Electrode:

Soln B.

a8 for Est Amnido Black

ub lor ¢3 6-BES Asiaho Bick

as for Est Amido Black,

incubated in the appropriate staining solutions

(Table 2),

Results

Chromosome studies; Fig. 1a shoks the karyo-

type of a female L. forresti, The diploid number

was 2N = 48. The largest chtomosome, desig-

nated pair 1, was acrocentric in IMVS§ 1, 2, 5,

7, 8 and 9. However, in IMVS 3. and 6, pair |

was heteromorphic, onc member hemg acro-

centric and the other being subacrocentric with

a distinct short arm. Pairs of 2 to 21 were

acToceninic forming a series graded in size,

Puirs 22 and 23 were small metacentrics. lhe

presumed X-chromosome was an acrocentric

representing about 6% of the total chromosome

length and the Y-chromosome an acrocentric,

The C-staining technique (Fig. 1b) showed

that although the centrometric area of some

chromosomes were C-banded, only in the small

Inetucentric pairs 22 and 23 was this marked,

In many preparations the presumed Y stained

slightly more intensely than other chromosomes

over its entire length.

Chromosomally Z. Jakedownensis differs

from L, forresti in possessing only one pair of

small metacenirics (Fig. 1c). Also, in the threc

individuals karyotyped, pair 1 was. always acro-

centric, Centromeric C-banding of L. Iake-

BIOCHEMICAL AND KARYOTYPIC STUDIES OF LEGGADINA LAKEDOWNENSIS 111

On An an

nn Aah Ha

oA AN AN

fin OO An

On AA AO

an an an

PA OAK Aw

on la An

hh Aa on Ag

AN AN fA OA

On AN 1& AO

an 10

ah &0

oh Of

Ah Ah AR an ly fe

10p 1c

mere

= it

quer

b% in | a!

& if 4 és =a

ee 23 ea -® 8

22 23

re—

10p 1b

' 4 a és 2

1 oe oe

> BQ

ge 8 8 ef Ae

*e “ff 88 Ae 88 4 5

2z 23 :

——

10p 1d xy

Fig. 1. Karyotype and C-banding of L. forresti and L. lakedownensis. 1a, Karyotype of female L. forrest

(IMVS 3) heteromorphic for pair 1;

for pair 1;

dawnensis (IMVS 12).

Species G-6-PD Hb TO

1b, C-banding of male L. forresti (IMVS 6) heteromorphic

lc, Karyotype of male L, lakedownensis (MVS 11);

1d, C-handing of male L. lake-

GA-3-PD Est-1 Alb NAD-MDH

Fig. 2. Representation of electrophoretic patterns observed for seven useful proteins in six pseudomyin

species. In each case the origin is to the left, and fastest migrating bands to the right (cathodal).

Key—L J, = Leggadina forresn; LA. = L. lakedownensis; Lid, = L. delicatula; L.h. = L. her-

mantsburgensis; P.n. — Pseudomys novachollandiac; P.a. = P. australis,

downensis (Fig. 1d) was evident only in the

smaller chromosomes. Tha presumed Y was

slightly more intensely C-bandcd than other

chromosomes over ifs entire length.

Electrophoresis: Of the 16 proteins studied

(Table 2), LAP, PHI, Tf, Est. 2 and Est. 3

showed evidence of polymorphism in at least

one species, and 6-PGD, LDH A & B and PGM

were identical for all species. This left seven

of the protcins studied that were consistent

within species but varied between species. The

electrophoretic results for these seven proteins

are shown in Fig. 2, and the resulting difference

matrix in Table 3

112 P. R. BAVERSTOCK, J. T. HOGARTH. S. COLE AND J. COVACEVICH

TABLE 3

Difference marrix for data in Figure 2

(Key a3 in Figure 2)

Lj, Lib Lid LA, Pn, Pea.

Ly, 0

Li 3 0

Td. 6 5 0

Lik, 6 5 a 0

P.n. 7 & 1 n 0

Pa 6 7 4 4 4 0

Discussion

Most L. forresti were found to possess the

same karyotype, although two were hetero-

morphic for a sub-acroventric pair 1, C-banding

showed that the short arm on the sub-acrocen-

tric member was not heterochromatic, suggest-

ing that the sub-acrocentric was telated to the

acrocentric by a pericentric inversion.

L, lakedawnensis, however, had a pair of

small metacentrics converted to a pair of acro-

centrics, presumably by a pericentric inversion.

Although a single fixed chromosomal difference

between L, forresti and L. lakedownensis is in-

suffizient in itself to indicate a species dif-

ference, taken in the context of the very low

karyotypic variation of the whole of the

pseudomyinae (unpublished data), a single

chromosomal rearrangement probably indicates

reasonable differentiation,

The biochemical data are mere convincing,

Of the seven uscful proteins studied, L. forresti

and L. lakedewnensis differ in 3 (Table 3),

This is considerable compared to the biochemi-

cal differentiation between 3 “good” species—

L. delicatula and L. novaehollandiae (1 dit-

ference), and £. novaehollandiae and L- her-

mannshurgensis (1 difference). These results

suggest that £. forresti and L. lakedownensis

may have been separated from each other for

at Ieast as long as have L. delicatula, L. navae-

hollandiae and L. hermansburgensis.

Phenetically L. lakedownensis and L. forresti

are biochemically more similar to cach other

thai either is to any other pseudomyin studied

(Table 3). Although more data are needed

these results support the maintenance of Legga-

dina aS a separate genus which at this time

would include only these two species.

Acknowledgments

We are grateful to Dr C. H. 8. Waits for

helpful comments during the course of this

study. The original specimens of L. lakedown-

ensiy were collected by R, Buckley and

presented to the Queensland Museum by C.

Tanner. We thank the various State Wildlife

authorities for permission to collect specimens,

several of which were collected by A. & J.

Robinson under an Australian Biological

Resourees Study Grant to C, H. S. Watts and

P. R. Baverstock.

References

ArricHt, F. E., & Hsu, T. C. (1971 ).—Localiza-

tion of heterochromatin in human chromo-

somes. Cytogenetics 10, 81-86.

Brewer, G. J. (1970)—“Introduction to Tsozyme

Techniques,” (Academic Press: New York

and London).

Hoimes, R. S., Coorer, D. W., & VANDEBERG,

J. L. (1973) —Marsuptal and monotreme

lactate dehydrogenase isozymes: phylogeny.

ontogeny and homology with eutherian mam-

mals. J, Exp. Zool. 184, 127-148.

SELANDER, R. K,, SMirH, M, H., Yano, &, Y.,

Jornnson, W. E., & Gentry, J. B, (1971).—

Biochemical polymorphism and systematics in

the genus Peromyscus. €. Variation in the old-

field mouse (Peromyscus pelionotus), Stud.

Genet. V1, 49-90 (Univ. Texas Publ. 7103).

Warts, C. H, S. (1976).—Leggadina lakedownen-

sis, a new species of murid rodent from north

Queensland. Trans. R: Sac. S. Aust. 100(2),

VOL. 100, PART 3 31 AUGUST 1976

TRANSACTIONS OF THE

ROYAL SOCIETY

OF SOUTH AUSTRALIA

INCORPORATED

CONTENTS

Foster, R. J., and Philip, G. M. Corystus dysasteroides, a Tertiary Holasteroid

Echinoid Formerly Known as Duncaniaster australiae - - 113

Crisp, M. D. Rediscovery of Acacia barattensis J. M. Black (Mimosaceae)

in South Australia - ~ - - - - - - i117

Mawson, Patricia M. Woodwardostrongylus obendorfi new species (Nematoda:

Amidostomatidae) from Kangaroos - - - - - 121

Callen, R. A., & Tedford, R. H. New Late Cainozoic Rock Units and Depo-

sitional Environments, Lake Frome Area, South Australia - 125

Glaessner, M. F. A New Genus of Late Precambrian Polychaete Worms from

South Australia - - - - - - - - - 169

PUBLISHED AND SOLD AT THE SOCIETY’S ROOMS

STATE LIBRARY BUILDING

NORTH TERRACE, ADELAIDE, S.A. 5000

CORYSTUS DYSASTEROIDES, A TERTIARY HOLASTEROID ECHINOID

FORMERLY KNOWN AS DUNCANIASTER AUSTRALIAE

BY R. J. FOSTER* AND G. M. PHILIP}

Summary

FOSTER, R. J., & PHILIP, G. M. (1976).- Corystus dysasteroides, a Tertiary holasteroid echinoid

formerly known as Duncaniaster australiae. Trans. R. Soc. S. Aust. 100(3), 113-116, 31 August

1976.

The type specimens of the nominal species Rhynchopygus dysasteroides Duncan 1877, Holaster

australiae Duncan 1877, Holaster difficilis Duncan 1887 and Galeraster australiae Cotteau 1890

(which include the type species of Corystus Pomel 1883, Galeraster Cotteau 1890 and

Duncaniaster Lambert 1896) are discussed and illustrated. All are included in one species correctly

designated Corystus dysasteroides (Duncan).

CORYSTUS DYSASTEROIDES, A TERTIARY HOLASTEROID ECHINOID

FORMERLY KNOWN AS DUNCANIASTER AUSTRALIAE

by R. J. Foster* and G. M. Putiiipt

Summary

Foster, R. J,, & Pitre, G. M, (1976).—Corystus dysastereides, a Tertiary holasteroid

echinoid formerly known a3 Duncaniaster australiae, Trans. R. Soc. S, Aust, 100(3),

113-116, 31 August 1976.

The type specimens of the mominal species Rhynchopygus dysasteroides Duncan 1877,

folaster australiae Duncan 1877, Holaster difficilis Duncan 1887 and Galeraster anstratiae

Cotteau 1890 (which include the type species of Corystus Pomel 1883, Galeraster Cotteau

1890 and Duncaniastér Lambert 1896) are discussed and illustrated. All are included in one

species correctly designated Corysius dysasteroides (Duncyn)-

Introduction

Holasteroid echinoids are not abundantly

represented in the diverse Tertiary echinoid

fauna of southern Australia, but there is one

common species'which. for the last eighty years

has been known as Duncaniaster australige

(Duncan). The purpose of this note is to

review the complex nomenclatural history of

the species and to decide on its correct

designation, Also, photographs of the type

material of four nominal species proposed by

Duncan (1877, 1887) and Cotteau (1890) are

published for the first time.

The species is known from the Tertiary

coastal basins of southern Australia from Eucla

Basin in the west to Torquay Embayment in

the east, and from New Zealand. The earliest

known Australian occurrence js in the Middle

or early Late Eocene; it is present in the Wilson

Bluff Lumestone al the Bluff and in Abrakurric

Cave, and in the Tortuchilla Limestone and

equivalents of the Si Vincent Basin, Jt makes

its last Australian appearance in the late Barly

Miocene (uppermost Longfordian) Watac-

poolan Limestone at Koonalunda in western

Victoria. The species also occurs in the South

Island of New Zealand; it appears first near the

base of the Weka Pass Limestone in the Early

Oligocene (questionable Whaingaroan), and

jast in the Gee Greensand in the Late Oligocene

or Early Miocene (Waitakian-Otuian). More

* BHP Oil and Gas Division, Melbourns, Vic, 3000,

stratigraphic details are given in a separate

paper (Foster and Philip, in press)-

Historical review

Dunean (1877, p, 49). described the species

Rhynchopygus dysasteroides from Castle Cove,

Victoria (Late Eotene Castle Cove Limestone)

and (1877, p, 31) described a further species,

HAfolaster australiae from the same jocality. The

holotype of R; dysasteroides is crushed, and it

Was ptesumably for this reason that Duncan

regarded the specimen as a cassiduloid. Pomel

(1883, p. 61) proposed the genus. Corystus for

R. dysasteroides because of its intercalary apical

system. In his revision of the Australian

echinoid fauna Duncan (1887, p. 421( pro-

vided a corrected woodcut of the apical system

of the holotype of H. auxfraliae. He recognised

that he had misinterpreted the species R. dysas-

teroides and been mistaken about its affinities,

As a consequence he renamed it Holaster

difficilis. Pomel'’s work was not well known at

the time and it is no doubt because of this

Duncan made no mention of the genus

Corystus.

Cotteau (1890, p, 548) described Goleraster

adustraliae from Mount Gambier (Early Mio-

cene Gambier Limestone) as a new genus and

species, placing the genus Galeraster close to

Holaster. Tate (1891, p, 276) first suggested

that A. difficilis and H. australiae were the same

species. In 1892 Bittner (p, 359) rejected the

t Department of Geoloxy and Geophysics, University of Sydney. N.S.W. 2006.

1t4

genus Cerysivs, noting Gregory’s (1890, p.

990) reference to 4. difficilis as an “unsatisfac-

tory species”, Also, in 1892 Tate published his.

strongly worded ¢riticism of Bittner's paper but

in regard fo these species he followed Bittner,

although he suggested that Galeraster australiac

oe an additional synonym of Holasrer ausira-

fae.

Lambert (1893, p. 97) transferred Al. aws-

iraliaeé 10 Pomel’s genus Lampadocorys but

later (1896, p. 317) made it the type species of

his new genus Duneaniaster which he placed

clase to Sregaster. Thus was created the widely

used name Duncaniaster australiae, In 1903

Lambert (p. 32) grouped the genus with Lam-

padocorys, Stegaster, Tholaster and Ojjaster in

his subfamily Echinocorynae.

Lambert & Thiery (1921, p. 332) recognised

Galeraster as a valid genus in the Bchino-

galerinae, stating (1924, p. 408) that Tate was

mistaken when he made Galeraster australiae a

synonym of Holaster australiae. They (1921, p.

364) reinstated the species Rhynchoprous

dysasteroides, and made Corysrus Pomel a

synonym of Rhyachepyeus dOrbieny, Last of

all (1924, p, 408), they relegated Duncaniaster

Lambert to a sub-genus of Cibaster Pomel.

H. L. Clark (1946), in his review “The

Echmoderm Fauna of Australia” mentioned

neither Corystus nor Galerasrer. He maintained

Duncaniaiter as a separate genus (p. 361}, but

did not consider it far removed from

Cardiaster; the only species he listed was D.

australiae (Duncan), Mortensen (1945, p, 84)

Tetained Cotteau's genus Galeraster in the

family Echinoncidae Wright and close to

Pyrina, but (p, 203) considered Corystus to be

a synonym of Cassidulus. He confirmed (1950,

Pp. 74) Duncaniaster in the Holasteridae close

to Cibaster. Wagner & Durham (1966, pp.

U445S U528) in the Treatise followed Morten-

sen in their placement of Galeraster and

Duncaniaster, and Corystus was tentatively

plated among the cassiduloids as a douhtful

nominal genus.

Type material

The holotype of Rhynchopygus dyavteraides

ts BM, E42418 (Fig. 2 C, B, F) and that of

Holaster australige is BM, E31067 (Fig. 2 A,

B, BD}. Both are lodged in the British Muscum

(Natural History), and both were collected

from the "No, 5 Upper Coralline Beds, Castle

Cove, near Cape Otway” in Victoria. This is

the old locality AWS of Wilkinson (1865) in

the Castle Cove Limestone, which Carter

R, J. FOSTER AND G. M. PHILIP

Fig. |. Plating of apical system of holotype of

Halaster austraiiae Duncan (BM £31067),

(1958, p, 21) refers to as his Foraminiferal

Units 2 and 3. The echinoids are probably from

the upper part of the formation in the latest

Late Eocene,

As indicated above, the type specimen of R-

dysasteroides is badly crushed, although the

adapical surface shows an holasteroid apical

system, similar to that of A. australiae (Fig,

1). In both spectmens the adoral surface is

poorly preserved. and the plastronal plating. is

obscure. Because of the state of preservation,

the presence or absence of a subanal fasciole

could not be established.

The holotype of Cottean’s Galeraster aus-

fraliae is an unnumbered specimen in the Ecole

des Mines, Paris, in the Cotteau Collection

(Fig. 2 G, H, 1), Its locality is “Mount Gam-

bier, Australia” and doubtless is from the

Gambier Limestone. The type section in the

sinkhole at Mt Gambier town is of Longfordian

(Barly Miocene) age, and Janjukian (Late

Ohgocene) outcrops are limited to restricted

areas NW and SW of the town. The precise

locality of Cotteau’s type, and of the only other

representatives of the genus from this forma-

tion {P20456 from the National Museum of

Victorian and T267a from the Tate Collection

labelled “Helaster woedsti Mt Gambier’), is

CORYSTUS DYSASTEROIDES, A TERTIARY HOLASTEROID

ECHINOID 115

Fig. 2. All natural size. A, B, D. Adapical, adoral, and lateral views of holotype of Holaster australiae

Duncan (BM 531067). C, &, F. Adoral, lateral and adap

ical views of holotype of Rhynchopy-

gus dysasteroides Duncan (BM E42418). G, H, I. Lateral, adapical and adoral views of holotype

of Galeraster australiae Cotteau.

1L6

nor known. The general echinoid fauna

presently available from the Gambier Lime-

stone appears to have its closest affinities with

that of the Longfordian Mannum Formation of

the River Murray cliffs In particular, 1267a

the only well-preserved specimen of Corystjis

from’) Mount Gambier was elsewhere {Foster &

Philip, in press) compared statistically with the

populations from a number of south-eastern

Australian localities ranging from Late Eocene

fo Early Miocene, and its parameters corre-

lated best with samples of populations from

the Mannum Formation and the Longfordjan

portion of the Port Vincent Limestone. It is

therefore concloded that the holotype is

probably from the Early Miocene. Again the

holotype is a poorly preserved specimen, It is

worn and cracked and a number of borings

occur in parts of the test. Sorface detail is

obscured by matrix and secondary calcite to the

degree that even the paths of the ambulacra are

difficult to trace, Preparation of the apical

region of the specimen showed the widely

separated oculars typical of an holasteroid

apical system,

R. J, FOSTER AND G. M, PHILIP

Conclusions

Despite the unsatisfactory nature of the type

matetial, We conclude that all specimens are

conspecific, We base this conclusion on the

large collections of the species available to us

from various localities in south-eastern Aus-

tralia. We here choose dysasteroides as the valid

mame for the species as it has page precedence

over qustraliae which was introduced by Dun-

can in the same publication, Pomel’s genus

Corystus has priority over Duncaniaster Lam-

bert. ‘Thus the valid Linnean species is Corysties

dysasteroides (Duncan).

In a further paper (Foster & Philip, in press}

we present a statistical analysis of samples of

Corystus populations ranging from Late

Eocene to Early Miocene in age. This analysis

is designed to depict the morphological trends

apparent in the evolution of the species. We

also have in preparation a taxonomic study of

all the holasteroid echinoids known from the

Tertiary rocks of Australia (including Western

Australia) and New Zealand. In this latter

article we will review the affinities of the genus

Corystus.

References

Brrrnun, A. (1892).—Ubher Echiniden des Tertiirs

von Australien. Sinz, A. Akad. Wiles. Wien,

(Math, Natur, Cl.) WI, 33371.

Carter. Av N, (1958),—Teniary Foraminifera

from the Aire District, Victoria. Bull. Geol.

Surv, Vict. 55, |-76.

Ciark, H. L. (1946),—The Echinoderm Fauna of

Aastralia, Its composition and origin, Publ,

Carnegie fast. Wash. 366, 1-567.

Correau, G. HL (1890).—Echinides notiveaux ou

pea connus, 2(9) Mem. Sov. Zool, France,

537-350,

Duncan, P.M, (1877)—On the Echinodermata of

the Australian Cainozoic (Tertiary) Deposits.

Quart. J, Géal, Sov. Lond, 33(1), 42-71.

Duncan, P. M. (1887),—A_ revision of the

Echinoidea from the Australian Tertiaries,

fhid. 43(3}, 411-430.

Fostra, R. J, & PHtur, G, M, fin press) —

Statistical analysis of the Tertiary holasteroicd

Corystuy dysaneroides from Australasia.

Thelassia Juwesl.

Grecory, 1. W, (1890),—Some additions to the

Australian Tertiary Echinoidea. Geol, Mav-

27, 481-492,

Lamperr, 3. (1893).—Etudes morphologiques sur

les plastron des spatangides. Bull. Sec, Sed,

hist, aot. ¥onne 47, 55-98.

Lamuert, |. (1896)—WNote sur quelques Echinides

crétaces de Madagascar. Bull. Soc. Geol.

Franoc 24, 313-332.

Camocrr, J. (1903).—Description des échinides

crétacés de la Belgique, J, Etude mono-

graphique sur le genre &chinocorys. Mém.

Mus, Roy, d'Hist, Nar, Belg, 2, 1-151,

Lamuerr, J., & Trtery, P. (1909-1925) —“Essai

de Nomenclature ralsonnée des Echinides” 1-9

{Libraire Ferriére: Chaumont).

Morrensen, T. (1948).—"A monograph of the

Echinoidea” IV (1) Holectypoida, Cassidu~

loida (C, A. Reitzel: Copenhagen),

MORTENSEN, T. (1950),—Idem V (1) Spatangoida

Power, N, A, (1883).—"Classification méthodique

et genera des échinides vivants et fossiles"

{ Adolphe Jourdan: Alger}.

Tate, R. (1891),—A bibliography and revised list

of the Described Echinoids of the Australian

Eocene, with descriptions of some new species.

Truns. Prac. R, Soc. 8. Aust, 14, 270-282,

Tate, R, (1892)—.Critical remarks on A. Biltner’s

“Eehiniden des ‘Tertiars von Australien”,

Sbid, 1§(2), 190-194.

Waonek, C, D. & Durnam, J. W, (1966).—

Holectypoids, Holasteroids.. Jn> Moore, R. C.

(Ed.) “Treatise on invertebrate palaeon-

tology” U3(2), 440-0450 & U523-U543.

Wirginson, C, S, (1865).—Report on the Cape

Otway Country (from Parl. Papers, 1864-

teal Repr. Geal, Surv, Vier, 1863-1864, 21-

REDISCOVERY OF ACACIA BARATTENSIS J. M. BLACK (MIMOSACEAE)

IN SOUTH AUSTRALIA

BY M. D. CRISP*

Summary

CRISP, M. D. (1976).-Rediscovery of Acacia barattensis J. M. Black (Mimosaceae) in

South Australia. Trans. R. Soc. S. Aust. 100(3), 117-120, 3 1 August 1976.

Acacia barattensis, previously known only from the type collection, has been rediscovered near the

type locality in the Flinders Ranges, South Australia. It is described in more detail than previously,

the legumes and seeds for the first time. Its taxonomic affinities and the type material are discussed.

Possible reasons for the species remaining "lost" for so long are suggested, and its state of

preservation is discussed in relation to its distribution and ecology.

REDISCOVERY OF ACACIA BARATTENSIS 5, M. BLACK (MIMOSACEAE)

IN SOUTH AUSTRALIA

by M. D. Crise*

Summary

Cer, M, D. (1976),—Kediscovery of Acacia baratrensis J. M, Black (Mimosaceae) in South

Australia. Trans, R. Sec. S. Aust. 100(3), 117-120, 31 August 1974,

Acacia barattensis, previously known only from the type collection, has been rediscovered

near the type lucalily m the Flinders Ranges, South Australia. It is described in moré detail

than previously, ihe legumes and seeds for the first time. Its taxonomic affinitics and the type

material are discussed. Possible reasons for the species remaining “lost” for so long are

suggested, and its state of preservation js discussed in relation to its distribution and ecology.

Introduction

Recently [ rediscovered Acacia barattensis

in the Flinders Ranges, South Australia. In so

doing [ established that the species was nol

extinct as previously supposed (Specht et al.

1974, p. 304), and that the locality informa-

tion given by its author was inaccurate. The

following formal taxonomic treatment? contains

new information about its morphology. type

material, affinities, ecology and distribution.

Taxonomy of Acacia barattensis

Acacia barattensis J. M, Black in Trans, R.

Sac, S. Aust, 56:42, t.1, fig, 2 (1932); Flor. 8.

Aust. ed. 2:419, fig. 578 (1948).

Somewhat diffuse, spreading shrub 2-3 m

tall with several slender branches arising at or

near the base, Breunchlets slender, glabrous,

gently curved so that the tips stand erect;

strongly angular and very viscid towards the

Ups; becoming terete, less viscid and faintly

striate towards the bases; greenish-brown but

usually covered with a black incrustation.

Foliage with a strong odour of resin acids

when dryy. Stipvles absent. Phiylledes erect,

natrow-linear but tapering slightly towards

beth ends, vertically flattencd, never terete,

gently incurved, abruptly rostrate-uncinate at

the upices, (3)5-8 cm long, 08-15 mm

broud, S-nerved on cach face, 1-nerved on

each margin (8-nerved in all); with a narrow

gtoove which is usually filled with brown resin

ahove and along each nerve; often shallowly

and irregularly sulcate between the nerves:

very sparsely and mihutcly pusticulate, initially

very viscid, Marginal gland scarcely visible,

2-3 mm above the base of the phyllode.

Peduncles 1-2 in the axils, each with a minute

narrow:triangular basal bract (0,5 mm long),

slender (0,2-0,3 mm diam,), terete, + papil-

lose-viscid, 8-13 mm long. Heads globular,

5-7 mm diam, at anthesis, ca 20-flowered

Floral bracts navicular with long (ca 0.5 mm)

triquetrous claws and with extended acute

apices, 0.8~-0.9 mm long and 0.2 mm bread,

densely papillose-viscid, the margins scarious,

+ entire, Flowery 4-merous, Calyx ca 1/3

length of corolla, consistently divided for 1/4

its length into triangular and barely acute lobes

with entire margins, papillose-viscid. Petals

oblong, recurved at the acute tips, 2 mm long,

faintly unimerved, + papillose-viscid, Joosely

connate for ca + their length. Ovary = sessile,

densely papillose-viscid. Leeurme narrow-linear,

coriaceous, straight or slightly curved, stipt-

tale, contracted to 4 its width between the

seeds, (6)8-10(15) em long, 2.5-3.5 mm

broad, often with a very narrow (1 mm) and

elongate (up ta 1 em) black sterile tip. Sur-

face of the legume initially very viscid, finally

not viscid, dark brown, with anastomosing

raised veins hetween the seeds, irregularly col-

liculate over the seeds. Margins of the legume

much thickened, straw coloured. Seeds longi-

tudinal, oblong-elliptic, smooth, dark brown,

ea 2.5 x 4.5 mm, Ari! much dilated. with

* Herbarium, Canberra Botanic Gardens P.O. Box 158, Canberra, A.C.T. 2601,

1 Personal communication: 1. Elix, Dept of Chemistry, Australian National University, A,C.T.

1i8 M. D. CRISP

Fig, 1. Acacia barattensis. A. Portion of branch in bud and flower. B. Phyllode. C. Transverse section of

phyllode, taken near the centre, showing resin-filled grooves. D. Apical portion of phyllode. E.

Inflorescence (note basal bract). F. Floral bract in lateral (left) and ventral (right) view. G.

Calyx and corolla. H. Legumes and peduncle. I. Exterior view of portion of legume over seed,

J. Seed, ari] and funicle. A-E from M. D. Crisp 889; F-G from M. D. Crisp 731; H-J from

M. D, Crisp 890.

REDISCOVERY OF ACACIA BARATTENSIS

hyaline wings, ance-bent above the seed and

gradually contracted through a sharp bend inte

a thrice-folded funicle.

Type citation; “Near Baratta head-station,

on a branch of the Siccus River and 20 miles

west of Kononamore.”

Holonpus: J. B. Cleland, 3.4i1,1930, “North

of Baratta Head Statn." (Raratta is at 31°59'S,

139°06E), AD 97338071 (f.)! fsorvpiz AD

96247254 (“belongs to AD 97338071") (41.)!,

Distribution; South Australia: Plinders

Ranges. Apparently restricted to the deep

gullics of an unnamed range forming the

northem and western boundaries of Biblianda

Station, ca 55 km E of Hawker and 16 km

NW of Baratta head station. All recent col-

lections have been made in the vicinity of the

peak known as “The Bluff".

Ecology; Apparently confined to oear-verti-

cal gorge walls, from just above creck level to

ca 30 m above, on skeletal soils between out-

cropping quartzite, Flowermg is apparently

irregular, having been observed in April,

October, November and December; fruiting in

October (immature) and November.

Additional specimens examined: Bibdliando

Stn. ca 2 km N of New West Bore (31°52'S,

139°OS' EE). M, D, Crisp T3L, 13.iv.1974. fi,

fr. & photo. (AD; CBG 060871—ortg, spec.);

Bibliando Stn, southern slopes of The Bluff,

West Bore Pudk (31°S1'S, [39°00 BE), M. D

Crisp 889, 20)%.1974, fl. & fr. (CBG 060873—

orig. spec., NSW); ibid., AZ, DB, Crisp 890,

20.x.1974, fi, & fr, (CANB, CBG 060872—

orig. spec., NSW, TL, US); Between Willipa

and Biblianiio, Af. G. Catford s.n., 2-xi.1974.,

ft. & fr. (AD 97448228—pro parted; ibid.,

M. G Carford som., 23.xi1.1974, fr, (AD

97448228—pro parle)

Affinities: In his original description. of

Acacia barattensix, Black placed it next to 4.

subporosa F. Muell. {the now scyregated 4,

cograta Domin). | consider it ta be much

closer to the group 4. gracilifolia Maiden et

Blakely, 4, wilhelmiana F. Muell., 4, helnisi-

ana Maiden and A. menzelii J, M. Black,

which apparently belongs in Bentham’s series

Calamifurmes. However it differs from all

these in its 4-mcrous flowers and in being

totally glabrous. Maiden & Blakely (1927) des-

cribe a 4-partite corolla for A. gracilifolia, tut

both their illustration and material examined

by myself hive a S5-partite corolla, In SE

Australia the most similar relative of A. barar-

119

fests is apparently A. merzelil, which has

stmilar phyllodes, glands, peduncles, legumes,

seeds and arils, However the latter differs in

having phyllodes shorter and usually terete, 2

fewer nerves per phyllode, bracts larger,

cucullate and prominently ribbed, flowers 5-

merous, floral bracts with short claws.

Discussion

The type materi] at AD consists of 2 sheets,

hoth annotated by Black. One of these (AD

97338071) hus two twigs of material and two

locality labels in Black's handwriting, Onc

label,

"North of Baratta

Head Siatn 3/12/30

(J. B. Cleland)

label marked "Kew 137°”,

appears to be the original information received

from Cleland, while the other,

“Baratla H.S. (on a branch of the Siccus

River & 20 miles W of Kaonamore.)”,

8 Black's transcription of the former for publi-

cation. There is no evidence to suggest that

the tWo twies are separate collections, Bath

twigs are identical in all respects, particularly

in the stage of flowering. The sheet also bears

extensive descriptive notes and drawings, and

a bold Jabel “Acacia Aaraztensis J. M. Black”,

all in his hand. Clearly this shect ts the holo-

type.

‘The second sheet (AD 96247254), origin-

ally kept separately in Herb. J. B. Cleland but

later transferred to AD, curries a locality label

similar to that on the holotvpe, and a twig

Wentical io the others, indicating that this

sheet is an isotype. The label is marked “Kew

2137" by an unknown writer, and

“Apparently a new sp., but if sent 1a Kew,

I scarcely know what to do—I.M_R,”

by Black. Cicarly it was after he wrote this

note that Black decided to describe: and name

the Acweia from the duplicate (holotype)

material, If any specimen had been sent pre-

viously to K. tt must have been returned,

because the only specimen of 4. buraltensix

now there is endorsed

“comm, J, M, Black, Jan 1933"

i.e. it Was sent there after publication of the

new species by Black, This third specimen

appears: in all respects to be a duplicate of the

holotype collection, and must be regarded as

A second isotype.

The ahove distussion shows clearly wiry

many attemprs to relocate Acacia haraitensix

falled. Whereas Cleland’s field locality was

120

“North of Baratta” (in the ranges where it

presently occurs and only about 10 km distant

from Baratta), Black’s published locality

erroneously focussed attention on the head

station itself, where it does not occur. The

populations of the Acacia are in fact restricted

to specific sites in the deep gorges of the

range at Bibliando. However, where popula-

tions do occur they are fairly extensive and

protected by the inaccessibility of their habitat.

Provided that no major disturbance occurs in

this range, Acacia barattensis is probably safe

from the threat of extinction for the immediate

future.

M. D. CRISP

Acknowledgments

I wish to thank the staff of the State Herbar-

ium of South Australia (AD) for the loan of

specimens and for assistance received. I am

also grateful to Dr Hj. Eichler of Herbarium

Australiense (CANB), who offered some use-

ful comments about the type material and

read the manuscript. Mr A. B. Court of the

Herbarium, Canberra Botanic Gardens (CBG)

offered many helpful suggestions. Dr A. Kanis

kindly located and supplied data from the

specimen at the Herbarium, Royal Botanic

Gardens, Kew (K).

References

MalweN, J, H., & BLAKELY, W. F. (1927).—Des-

criptions of fifteen new Acacias. J. Roy. Soc.

N.S.W. 60, 171-196.

Srecur, R. L., Ror, E. M., & BouGcuron, V. H.

(ed.) (1974)—Conservation of major plant

communities in Australia and Papua New

Guinea, Aust. J. Bot, Supp. Ser. 7.

WOODWARDOSTRONGYLUS OBENDORFI NEW SPECIES

(NEMATODA: AMIDOSTOMATIDAE) FROM KANGAROOS

BY PATRICIA M. MAWSON*

Summary

MAWSON, P. M. (1976).-Woodwardostrongylus obendorfi new species (Nematoda:

Amidostromatidae) from Kangaroos. Trans. R. Soc. S. Aust. 100(3), 121-123, 3 1 August 1976.

Woodwardostrongylus obendorfi n.sp. is described from the oesophagus of Macropus parryi

(type host), M. robustus, and M. rufogriseus. It is distinguished from W. woodwardi (Wood) chiefly

in having only 6 pairs of oral denticles instead of 16. Woodwardostrongylus Wahid is transferred to

the family Amidostomatidae, and the genus Cristaceps Mawson is placed as a synonym of

Woodwardostrongylus.

WCODWARDOSTRONGYLUS OBENDORF! NEW SPECIES (NEMATODA;

AMIDOSTOMATIDAE) FROM KANGAROOS

by Parricia M. Mawson*

Summary

Mawson, P. M. (1976).—Woodwardostrongylus obeadorfi new species (Nematoda: Amido-

stromatidae) from Kangaroos, Trats. R. Soe. S. Aust, 100(3), 121-123, 31 August 1976.

Woodwardostrongylus obendorfi n.sp. is described from the oesophagus of Macrepus

parryi (type host), M. robastus, and M. rufogriseus. It is distinguished from W. woodwaurdi

(Wood) chiefly in having only 6 pairs of oral denticles instead of 16, W’oodwardostrongylus

Wahid ts transferred to the family Amidostomatidae, and the genus Cristaceps Mawson is

placed as a synonym of Woodwardostrongylus.

Introduction

The genus Woodwardostrongylus was

erected by Wahid (1964, p. 184) for Pharyn-

gostrongylus woodwardi Wood, 1931, Mawson

(1971, p.. 174) not having seen Wahid's work,

proposed the genus Cristaceps for the same

species, pointing out that this genus was close

to Filarineme in the family Amidastomatidae

(serisu Inglis 1968). Cristaceps now falls as a

synonym of Woodwardostrongylus, but the

latier must be transferred to Amidostomatidae.

Nematodes recently taken from the veso-

phagus of three species of macropods have

heen identified as a new species of Wood-

wardosirongylus, In all three cases the worms.

were threaded through the oesophageal epi-

thelium so that care was needed to collect

them entire. This situation is similar, though

in the oesophagus instead of the stomach, to

that occupied by W. woodwardi in the two

recorded findings (Wood 1931; Mawson

1971). It is a locale which is likely to escape

all but the most careful dissections, so it is

possible that species of the genus are more

widely distributed than the records indicate.

T am very grateful to Dr Brian Coman and

Dr Tom Kirkpatrick who shot the kangaroos

and to Mr David Obendorf who first noticed

the presence of the worm.

The micrographs (Figs 10, 11) were taken

by E.T.E.C. Avtosean in the Central Electron

Optical Laboratory of the University of

Adelaide, I am indebted to Dr Karl Bartusek

of this Laboratory for help in taking the micro-

graphs, and to P. G. Kempster for developing

and printing them.

Woodwardostrongylus obeadorfi n sp.

FIGS 1-11

Host and Locality: Macropus parryi (Bennett)

(type host and M, rebustus (Gould), fram

Dorrigo, N.S.W.; M. rufogriseus (Desmarest}

from Warwick. Qld,

‘The wortns are thin and clongate, the males

15,9-16.7 mm, the females 24-26 mm in

length. The. body, especially of the female, is

widest in its posterior part. The rounded

anterior end bears a small round mouth, on

each side of which lie six prominent denticles

each associated with a plate-like sclerotisation

in the cuticle. The mouth leads to a thick-

walled buccal capsule or vestibule. The lumen

of this is narrow but wider dorsoventrally than

from side io side. The walls are faintly striated

transversely (more: distinctly in some specimens

than others), and are distinctly thicker pos-

teriorly than anteriorly. The cephalic papillac

and amphids are very small.

The oesophagus widens in its posterior half

to a very slight terminal swelling. It is 800—

900 ym long in the male, 900-1050 «xm in the

female. In the male the distance from the

anterior end of the worm to the nerve ring ts

320-400 pm, to the cervical papillae 300—

460 wm, and to the excretory pore 440-510

wm; in the Female these distances are respec-

* Department of Zoology, University of Adelaide, Adelaide, S. Aust. 5000.

122

PATRICIA M. MAWSON

200 pm

Figs 1,2 and 3—Head, in semi-cn face, lateral and ventral views respectively. Fig, 4—Oesophageal

région. Figs 5, 6, 7, and 8—Views of bursa. Fig, 9—Posterior end of female. Figs 1-3 to same

scale. Figs 5-8 to same scale.

tively 360-400 pm, 560-600 pm, and 500-

550 pm.

The bursa is only slightly lobed, closed

ventrally, and somewhat voluminous dorsally

where it extends so that the dorsal ray for

most of its length lies at right angles to the

long axis of the body. The arrangement of the

rays is shown in Figs 5-8. The genital pore is

of medium size, apparently without accessory

lobes. The spicules are 1700-2100 ym long,

the ratio body length: spicule length being

9.9-12.8. A gubernaculum is present.

In the female the tail is 180-220 ,m long,

conical ahd pointed. The vulva is shortly in

front of the anus, 300-350 «m from the pos-

terior end, The vagina is relatively long, up to

800 ym. Vaginal eggs measure 140-150 x 70-

80 wm.

The species is distinguished from W. wooa-

ward? mainly by the presence of only six pairs

of oral denticles instead of sixteen pairs, and

by the presence of the associated basal plates,

which are not seen in the type species. There

is also a difference in the site in which the

species occur in the body, W. woedwardi in

the stomach and the new species in the oeso-

phagus. In Md. parryi and M. rufogriyeus the

worms were numerous, but only one was

found in M, rebustus.

WOODWARDOSTRONGYLUS OBENDORFI FROM KANGAROOS 1

Fig. 10. S.E. Micrograph, anterior end almost en face (x 1500).

Fig. 11. S.E. Micrograph, part of region around mouth, showing two of the submedian papillae, the

mouth, and some of the oral denticles (x 3500).

References

Inciis, W. G. (1968).—The geographical and Wann. S. (1964).—A preliminary revision of the

evolutionary relationships of Australian genus Pharyngostrongylus Yorke and Maple-

trichostrongyloid parasites and their hosts. J. stone, 1926. J. Helminthol 38, 180-190.

Linn. Soc. Lond. Zool, 47, 327-347.

Mawson, P. M, (1971).—Pearson Island Expedi- Woop, W. A. (1931).—Some new parasitic nema-

tion 1969.—8 Helminths. Trans. R. Sac. S. todes from Western Australia. Rep. Director

Aust, 95, 169-183. Inst. Animal Path. Univ. Camb. 1, 209-219,

NEW LATE CAINOZOIC ROCK UNITS AND DEPOSITIONAL

ENVIRONMENTS, LAKE FROME AREA, SOUTH AUSTRALIA

BY R. A. CALLEN* AND R. H. TEDFORDT

Summary

CALLEN, R. A., & TEDFORD, R. H. (1976).-New late Cainozoic rock units and depositional

environments, Lake Frome area, South Australia. Trans. R. Soc. S. Aust. 100(3), 125-167, 31

August 1976.

Five new rock units are defined for the Lake Frome area of South Australia.

The Namba Formation of Miocene age constitutes fine grained immature muddy sediments laid

down in a low-energy fluviatile and lacustrine environment, possibly partly estuarine or lagoonal.

Climate was subtropical or warm temperate with high rainfall, but seasonal aridity. Aphanitic

oolitic lacustrine dolomite and palygorskite are included in this sequence. The Flinders Ranges had

very low relief. The overlying and intertonguing Willawortina Formation represents alluvial fan

deposits with minor lacustrine phases, recording the beginning of the late Cainozoic uplift of the

Flinders Ranges, during which the Miocene lake was greatly reduced in area.

The Millyera Formation, constituting laminated ostracode bearing clay, fine sand, and charophyte

limestone, records lacustrine deposition during the Pleistocene. This took place in an enlarged

ancestral Lake Frome. The essentially fluviatile and aeolian deposits of the Eurinilla Formation and

Coonarbine Formation were deposited during the late Pleistocene and early Recent. Arid and

pluvial climates alternate in the late Tertiary and Quaternary. Drainage trends and the predecessor

of Lake Frome were established, closely approximating present day geography. During deposition

of the Coonarbine Formation the seif dunes of the southern Strzelecki Desert formed.

NEW LATE CAINOZOIC ROCK UNITS AND DEPOSITIONAL ENVIRONMENTS,

LAKE FROME AREA, SOUTH AUSTRALIA

by R. A, CALLEN* and R. H. TepFoarp{

Summary

CALLEN, R. A., & Teprorp, R. H. (1976) —New late Cainozoic rock units and depositional

environments, Lake Frome area, South Australia. Trans. R. Svc, S. Aust, 100(3), 125-167,

31 August 1976.

Five new rock nnits are defined for the Lake Frome area of South Australia,

_ The Namba Formation of Miocene age constitutes fine grained immature muddy sediments

taid down in a low-energy fluviatile and lacustrine environment, possibly partly estuarine or

lagoonal. Climate was subtropical or warm temperate with high rainfall, but seasonal aridity.

Aphanitic oolitic lacustrine dolomite and palygorskite are included in this sequence. The

Flinders Ranges had yery low relief. The overlying and intertonguing Willawortina Forma-

tion represents alluvial fan deposits with miner lacustrine phases, recording thé beginning of

the late Cainozoic uplift of the Flinders Ranges, during which the Miocene lake was greatly

reduced in area,

The Millyera Formation, constituting laminated astracode bearing clay, fine sand, and

charophyte limestone, records lacustrine deposition dying the Pleistocene. This took place in

un enlarged ancestral Lake Frome. The essentialfy fuviatile and aeolian deposits of the

Eurinilla Formation and Coonarbine Formation were deposited during the late Pleistocene

and early Recent, Arid and pluvial climates alternate in the late Tertiary and Quaternary.

Drainage trends and the predecessor of Lake Frome were established, closely approximating

present day geography. During deposition of the Coonarbine Formation the seif dunes of the

southern Strzclecki Desert formed,

Introduction

Mapping on the FROME (Callen 1975},

and CURNAMONA 1:250000 geological

sheets has resulted in differentiation of several

Tertiary and Quaternary rock units which can

be traced throughout the Lake Frome area

(the region south of Lake Callabonna between

the Flinders, Barrier and Olary Ranges). The

Eyre Formation has been detined previously

CWopiner et af. 1974). It lies immediately

beneuth the units described here for the Lake

Frome area and can be recognised over a

mucit wider region. The other units are at

present restricted to the Lake Frome region,

though correlation with units elsewhere,

especially in thy Lake Eyre Basin, is generally

possible on a firm basis.

There was a low divide between the deposi-

tional areas of Lakes Frome and Eyre, sug-

gested by the distribution of arenaceous

material in the Miocene rocks. The develop-

ment of this divide is clearly described by

Wopfner (1974, p, 6). Thus the Lake Eyre

and Lake Frome areas formed two distinct

depositional basins during late Tertiary times:

different sets of formal names are used for

rock units in each. In late Tertiary and

Quaternary times the Flinders Ranges, achieved

their present dimensions, completely separating

the two basins by a range of mountains.

This paper describes five rock units requiring

formalization under the Australian Code of

Stratigraphic Nomenclature (1973), com-

menting on the paleo-environmental inferences

to be drawn from them. The nomenclature

supersedes that shown on the FROME. geo-

logical map, relationships between units. now

being on firmer basis. The paper is divided

into: two parts, dealing with essentially Tertiary

and Quaternary units respectively. New geo-

graphic names have been formalized with the

Geographic Names Board of South Australiz

* Gealogical Survey of South Australia, Box 151 Eastwood, S. Aust, 5063.

t American Museum of Natural History, Central Park West, New York, U.S.A. 10024_

126

(pers. comm. 1973) and are designated with a

superscript wherever they first appear, thus:

Lake Namba*. Geologic names have been

cleared with the Central Registry (Canberra,

1973). The paper derived from a report by

Callen (1974)3 and an MLSc. thesis (Callen.

1976) 4. Additional stratigraphic data may be

found in this thesis.

Previous work includes the early geological

surveys of Selwyn (1360), Brown (1884) and

later of Jack (1930) and Kenny (1934), More

recently Ker (1966), Krinsley er al. (1968)

and Draper & Jensen (1975, in prep.) have

reported on hydrology and geology. The mar-

gins of the basin have been the subject of

regional mapping programmes by the South

Australian and New South Wales geological

surveys, on 1:250000 scale. Relevant to this

report are Leeson (1967)3, Firman (197134

and Coats (1973). A detailed basin study of

the older unit NAMBA FORMATION is in

progress and will be reported at a later date.

A preliminary account of the stratigraphy 3s

presented in Callen (1976), which gives the

structural and tectonic setting,

The terms used ta describe the sedimentary

rocks are those of Folk er af, (1970), ynless

indicated otherwise, Colours are given sym-

bolicallv in terms of Munsell Colour Code

(Geological Society of America 1951). A

relative scale was used for designating the

thickness of cross-bedding, as follows: very

small <J] cm, small 1-5 cm. medium 3-50 cm,

large 0.52 m, very large >2 m- In the

designation of contact features, core width

plices a limit on the interpretation, as it does

on maximum grain size: cobbles and boulders

are interpreted from the proportion and shape

of fragments ground down and broken by the

drilling operation, and nature of petrophysical

log response,

The older units (Pt i) were described mainly

from bores, the younger (Pt 11) from outcrop

Knowledge of the younger units was derived

trom detuiled investigation of over 100

trenched enlcrop sections, Where passible

units were raced between sections, Fossil soils

were an aid lo stratigraphic interpretation.

R. A. CALLEN & R. H. TEDFORD

The location of the sections is shown in

Fig. 1, Tables | and 3 summarizing rock unit

properties, palacontology and gtomorphology,

Svmbhols are in Fig, 2,

The subsurface sections were studied from

cores derived from bores drilled by the South

Australan Department of Mines and private

companies, Some percussion and rotary curting

were used to assist correlations, hut those

utilized for type sections were cored continu-

ously, and are availuble for inspection at the

South Australian Department of Mines Core

Laboratory, Petrophysical logs were run in all

cases, The lithological descriptions were sup-

plemented by binecular microscope exanina-

tion, and clay (x-ray diffraction) and grain

size (sieve and pipette) analyses were per-

Formed by Drs R, N. Brown and B, G

Stevenson respectively of the Australian

Mineral Develapment Laboratories,

The text is regarded as a supplement to the

diagrams and tables, descriptions in Tables 1

and 3 should be read first. Complete descrip-

tions of each scction are given in the appen-

dices, wherein the sequences are described as

they occur on the earth's surface—i.e.

youngest at top, oldest at base. Depths to the

top of cach unit or bed from the bore collar

are given. and the thickness is placed at the

start of its description, Jn cach unit. descrip-

tion of the dominant lithology ts capitalized;

followed by qualifying descnoptors referring to

each lithology in the same order.

Division into units in the reference scctions

is Intended as sn aid to identification of the

appropriate intervals in the descriptions (Scec-

tion 12, Fig. 3), fet a formal subdivision,

Core loss is indicated in the hore Ings (Fig, 3

sections 10, 1) & 12).

Pt. |\—Older Cainozoic Nock Units

A weneral definition of cach unit giving

satient Eealures, age and gcamorphic setting is

presented in Table 1, representative sections

in Fig. 3. Appendix | gives detailed desetip-

tions of individual anets

i Callen, R, A, (1974)—New Rock Units and Climate of the Cainozoic, Lake Frome area, South Auis-

tralia. §, Aust. Dept. Mines Rept. 74/75 unpub.

+Oallen. K, A. (1976).—Stratigraphy, sedimentology and uranium deposits of Ternary rocks, Lake

Frome srea, South Australia. M.Sc. thesis, University of Adelaide (unpublished)-

* Leeson, B (1967),—Geology of Bulcanoona 1:63 360 map area. 5. Aust, Dept Miiey Repl. RB 64/92

uligub,

‘Firman, 7, B, (1971) —Regional stratigraphy of surficial deposits in the Great Artesian Basin anit

Frome Embayment in South Australia. S. dws. Dept Mines Rept. RB 71/06 unpub.

CAINOZOIC ROCK UNITS 127

al Gs 4 = ee

| MARREE J, 5 fe CALLABONNA

2 BLANCHE =f

SCALE IN RILOMETRES

io ac} 19 zo 30 4c 40

rhs, g MINAD LEUZ BORE

TAKE

CALLABONNA

Lute Yeurieros fy betes bes I

Late drag rirdery

STREZELECRI

thake Sim denen Jo ah

“ . '

wou (untnariow Cont! Gaye rbenf

‘ i

4 By ii

’ Pre

bebe Cooaer

\ a bite Hundetan

. Fade Cuesierione

y - _ oF ai Qi bets Bumtardon

i See IN ys “ .

fe aa WOOLTANA'L BORE @ ye ty ‘i DESERT

PONE SN UNS tO Paes ni (ft Foie

Ree: Wareinony” i tate ovary Te

a w Hi

E i ~ Late Mallanitpen Y

oy : Take Pulorrs |

NY PAURALIO HORE, 34)

Tube Karp, 2 of

atinke Gonutary

etude Of dub Biman se

Wa eS ee

“ew ardvambe'

4 Avena)

Late Anke

. 1 1 <2) anes

as Og a x 1 2 | SENS Eo

& af od ‘ean 5 OORE / vas ghute Yomtanein | . JAI, Wistamysieelea

frome Dees! a Lave tHrkarnatas aia alae : he

a Sir Shute Moyet Mf >

% sf il vas alae | on

2\.

5 |

} +

)

fhti'iurasive

LEGEND

TAINOFOIC # MESGIOR ___ oad

He Cretateons

ancien ove Preccmbero ced mune

weal Ceysteliee basemen!

bade Eyre '

Hl Hinrent

LOGALITY Deane

" \

| Lal

( Ligke Torrens.

ou.

ULE ARSED

Wh of

OO ee az

reser! ben tvel Meineny selina —— oe

Edqeonl hiya leeal olin -__aeae

410

lyezotartipr sone Wet vwrhas oe

ren

leg tem 2 __ in

KILOMETRES

¥ Soo 400, MERC HIE

LOCALITY MAP 2A civeh

FA Colla

KILOMETRES

a 1000

Hrente bet) — ————— ee

Te EM Rope

Fig, 1, basally map, Lake Frome arca—location of type sections. Numbered sections shown in Figs 3,

" '

128 R, A. CALLEN & R. H. TEDFORD

LEGEND

LITHOLOGY SEDIMENTARY STRUCTURES

Cobbles, boulders S | Horizontally lominated

Pebbles Cross laminated

Very small scale (sets <lcm thick)

Very coarse grained sand—granules

vy | Small scale (sets lem--Sem thick)

Very fine grained—coarse grained sand

“o— | Medium scole {sets Scm—O.5m thick)

Silt

L Large scole Isets >0.5m thick)

7 :

arth | Burrows, burrowed contact, trace fossils

Bedding planes

Flot/wavy — Sharp vedding plone

TH Flat/wavy — Transition <lem

Intrafarmational clasts

Sandy limestone: (or dolomite)

@ BD) Silcrete, limestone clasts

Carbonate nodules (primary)

FOSSILS

Charophytes

Spores and pollen

Primary gypsum

Mica

Calites, pisolites

SECONDARY ALTERATION

Algal stems

Vertebrates (other than tish|—aqualic or terrestrial

24 Manganese staining

Fish bones

Secondary calcareous cement of

groundwater calcrete

Gastropads

Ferruginous or manganiferous matiling

{"matmorisation”)

Land snails

Calcareous paleosols '

(length indicates degree ot development)

Ostracods

Aboriginal artifacts

Emu egg shell

CLAY MINERALOGY

S = smeclite

Rl = randomly interstratitied cloys

M = mica/illile

K = kaolinite

ELECTRIC LOGS

SP = self potential

— Loyer of ailphide or iron and mangenese oxide R = point resistivily

74-1063 Del. 7S. R.A. Catlen S.A, Dept, of Mines

Fig. 2. Legend.

CAINOZOIC ROCK UNITS

LITHOLOGY

NAMBA FORMATION § (Derivation, Lake

Namba*?, CURNAMONA map sheet. “Namba”

is the Jadliauray aborigin! tribal word for bone-

fish).

The type section is Yalkalpo No, 1 bore

(section 12, Fig. 3) drilled by the South

Australian Department of Mines, Though not

typical in some respects, this is the only section

demonstrating the relationship to the Eyre

Formation (Appendix 1). The sequence is of

reduced thickness (56.60 m} compared with

that of the reference section in Wooltana No,

1 bore (170.09 m, Section 11, and Appendix

}, excluding unit 4 which is 68.24 m thick),

Section |! also contains 4 microflora important

in the age determination of the unit, The most

extensive outcrops are on the west shore of

Lake Tarkarooloot {Fig 14), where 26 m

are exposed, and at the south and north ends

of Lake Namba (e.g Section §, Fig. 15).

These are unsuitable for designation as type

sections, as only the uppermost heds are

represented.

The type section consists of a series of

eyelic sand/clay sequences in the lower part

totalling 32 m of mterhedded yellowish silt

and dark grey or olive clay. This ts overlain

by 24.8 m of burrowed yellowish silt and

intraformationally brecciated light olive clay.

The cyclic sequences constitute the following,

from the base of cach set upwards:

fa) Fine to medium-grained sand with small

to medium scale cross stratification,

Laminated silt to very fine sand with very

small scale cross-lamination.

A zone of dolomite or caleite patches or

a bed of dolomite.

A relatively thick dark grey clay with

uregular shiny-surfuced fractures [skew

planes of Brewer 1964} and scattered

patches and wisps of fine to coarse sands,

in Which grains are polished,

In this sequence (a) and (b} may alternate,

or either (a) or (b) may be absent. Unit 5 of

the type section represents a relatively com-

plete evele;

170m Altemating CLAY and SILT to SAND.

Sand very fine grained moderately sorted.

percentage increasing upwards. Grains

very angular, with crystal faces developed

on quartz. Beconles calcareous al top.

Bedding lenticular, with sedimentary

brecciation and = possible burrowing

aclivity. Obscure horizontal lamination at

top. 7% carbonate grains, rare mica,

Colowr 5Y6/1 mottled lOYR6*6.

(ob)

tc)

fd)

129

W.75m SANDY CLAY. Vertically streaked trai

sition zone from sand to clay. Fine sand

forms streaks and patches in clay. Very

poorly sorted medium silt, with modes in

clay and very fine sand sizes,

CLAY, black (SY1/1) and tough, with

characteristic frregular shiny-surfaced

fractures, and streaks af white carbonate.

Matted with orange brown colours which

SugeesE an irregular microstructure. Many

brown patches have well defined straight

boundaries, producing angular blocks

with dendritic or patchy internal struc-

jure, Unoxiclized clay in these blocks is

ereenish grey. Scattered patches of silt

and very fine sand are present, Upper

contact sharp, but disturbed, with partial

mixing into overlying sand,

The hurrowed silt is very finely laminated,

but this is often disrupted by burrowing, Very

fine sand sized maternal is the coarsest grain

size encountered. Colours are maiuly yellowish

grey to yellowish-white for silt or light olive

for clays, having a greasy lustre. Fractures do

not reach the degree of development of com-

parable structures in the black cluys of the

lower part of the sequence.

4.10 m

The burrow structures are « few millimetres

in diameter, containing convex-down Jamellae

usually Jess than 0.5 mm thick. They are

irregular and often branch, tending to be con-

centrated in certain hortzons. Many of the

homogenized clays have a churned structure

suggestive of bioturbation,

The top of unit 9 marks the last appearance

of the tough black clays characteristic of the

lower part of the formalion (Fig, 4}, Pre-

quently alunite (KAl4(SO,).(OH),) is

developed as lustrous white particles or patches

within the clay at the top of this unit. Above

unit 9. silts dominate over clay, and burrows

(Fig. 9). are more common.

The outcrop at Lake Tarksrooloo (section

13, Fig, 14) as situated on the western cliff

face, immediately north of the track-crossing,

on the route from “Frome Downs” to Black

Oak Bore, The lower part of the section is a

few tens of metres south of this track. The

two parts were corfelated Using continuously

traced bedding planes, and levelled with an

Abnocy hand level. The strata ure essentially

horizontul, as are those in the type section.

Nolable features of this outcrop afe the

interbedded gypsum nodules in the upper part

of the seclion, the presence of ostracode-

bearing aolitic dolomite associated with paly-

gorskite, burrowed fine sand beneath the upper

clay-dolomite sequence, the finely Jaminated

130

calcareous silt near the base of the sequence,

and the sharp contact with the upper tough

black clay, ‘These features, particularly the

last mentioned, are useful in correlation. In

section 12, the petrophysical logs indicate ihe

interval between units 12 and 13, which lacks

core, is probably silcrete, calcrete or dolomite,

The absence of palygorskite beneath it suggests

it may not be dolomite (this clay mineral is

invariably associated with dolomite elsewhere

in the basin. A turtle shell fragment in unit

10 supports a lithological correlation with

section §, if the black clay und ?dolomite are

also correlated, but this is not in agreement

R. A, CALLEN & KR H. TEDFORD

with the clay mineralogy. Section 13 shows the

typical dolomits—palygorskite association, and

trend towards illite deamination tn member two.

Typical of the Namba Formation outcrop

ate the brown chert nodules which cover the

breakaway slopes, and black manganese oxide

comling on the grains in sand heds Micro-

scapically, the chert sodules have structures

indicative of shrinkage and formation from

accretionary silica gel. The black stain as man-

ganése, Both these secondary effects are locul-

ized, occurring in sands cropping out in the

banks of stream valleys eroded in the Namba

Formation, prior to the deposition of the

Figs 4-9. Older units, Examples of Namba Formation lithology. Scales in mm and cm, Core sections.

Arbows point 10 top of section.

Fiz, 4. Section 12, Yalkaipo | bare, 125.00 m. Core, Dark grey clay with streaks of carbonate,

darker and lighter clay, and sand, some filling burrows or root holes. Vertical disposition of

patches Well-displayed, Kepresents swamp deposition or a lake deposit which has been subject

10 Ssubadrial exposure. Centripelal orientation of streaks is result of expansion of clay as it

enters the cuore barrel,

Section 3), Wooltarta 1 bore, 218.68 m. Section through core, showing upper contact of

laminated dolomite bed, Shrinkage and cracking of the dolomite has occurred, allowing pene-

tration Of the semi-fluld overlying clayey lime (CJ. Represents chemical sedimentation in a

lacustrine or marginal marine environment. Boundsries of carbonate fraements and laminae

. Section [1, Wooltanu 1 bore, $8.72 m. Core. Calearcous claystane with numerous burrdws in-

Alled with green-grey clay, Irregular shrinkage crack (C) has been infilled with semiliquid

clay which carrics carbonate particles. The clay-filled crack is itself burrowed. indicating

genesis soon after deposition, Represents combined chemical and detrital deposition in a mar-

. Section 11, Wooltana 1 hore, 122.00 m. Core. Fine lamination with typical alternation of sili

and sand, Very fine scale ough cross-lamination.. Quiel water deposition (migrating cipples!

. Wertaloana 1 bore, 142.00 m. Araldite peel of sectioned core, Typical example of small scale

cross-lamination in medium grained sand, partly disrupted by burrowing in upper part. Clayey

laminae alternate at base, Relief? coincides with porosity, though affected by varying thickness

of core acrass section, Cross bedding formed by ripple migration, in un offshdre bar or

. Section 12, Yalkalpo 1 bore, 22.60 m, Core. Top of bedding plane, Shows burrows along bed-

ding plane, with concave internal lamination (0). Represents quiet water depesition with bur-

(0-13. Oulcrop of Numba Formation and Willawortina Formatien, core of Willawortinn Fotma-

10 Vertebrate fossil float from the Namba Formation of L. Yanda on Eugigilla Créek. Vertebra on

far left (D> is riverine dolphin, on its right (L) are tungtish teeth and two fish spines (F}

Jates. A Cragment of bird bone (B) is

- South end of Lake Namba. Typical outcrop of Namba Formation. Gypsum nodule cappsing

(G) overlies thin nodular dolomite (white: LS), Greyish olive silty clay (grey) occupies. most

of section, Gragsed white beach at base of slope is very fine grained Iaminated sand (8), 30

em scale rests on upper contact in trench, Outcrop surface is covered hy gypsum nudules and

dlomite probably

. Baleanoona Ck, Willawortina Formation. Calcified mediwr ¢rossbedded sand Jens in cule

cxreous reddish brown very poorly sorted clay-sill. Note thin bedding in silt, Sand lens repre-

sents deposition from: higher powered streams, fine sediments are floodplain deposits, Scale 30

Fiz. 5

emphasized by inking,

Fiz. 6

ginal Marine Jagoon or lacustrine environment, with burrowing organisms.

Fig. 7

in o tidal, lacustrine or floodplain environment,

Fig 8

channel,

Fig ¥

fOowinle Organisms,

Figs :

Hon,

Fig.

In centre (F,C) are mainly crocodile scutes and turile r

on upper right corner, From base of upper unit of Namba Formation, Scale 30 cm,

Fig. 1

weathered clay, Sand represents channel of tloodplain deposition, clay and

Jacustrine.

Fig. 12

cm.

Fig. 13

. Section 10, WC2 bore, 68.75 m, section of core. Willawortina Formation shows large pebbles,

granules, very coarse silty and clayey sand. Extremely poor sorting. Represents deposition in

an alluvial fan environment, Scate in mm.

CAINOZOIC ROCK UNITS

Neoned

Hem vera

Mow

132

Millyera Formation and Eurinilla Formation

(new names see Pt. II).

Wooltana No. 1 bore (Fig. 3, Section 11

and Appendix 1), drilled by the Australian

Department of Mines is an important supple-

mentary section, exhibiting a thicker sequence,

lithologically more typical of the Namba For-

mation than the type section, It also demon-

strates the intertonguing relationship with the

Willawortina Formation (new name Pt. 1).

The base of the Namba Formation was not

penetrated, though cuttings from old Pootana

bore (Fig. 1, 50 km north-north-east of Wool-

tana No, 1 bore) indicate a total thickness of

190 m. This compares with 54.40 m in Yal-

kalpo No. 1 bore (Section 12). The sediments

have been divided into six informal units. The

lowest of these (unit 1) consists of 8.5 m of

laminated black and dark olive carbonaceous

clays with characteristic fauna and microflora

(discussed later), Laminae containing ostra-

codes of early Neogene aspect (including

cypridids—pers. comm. K. McKenzie 1973),

and fish spines are present. Protoconchs of a

small gastropod (Potamopyrgus s.1., see Lud-

brook 1972)5, are scattered through the clay

and ?gastropod tracks and burrows of other

organisms are common on_ bedding planes.

These sediments are restricted to the Poontana

Sub Basin west of Lake Frome.

Unit 2 (40 m) is dominated by white, fre-

quently oolitic, dolomite beds (Fig. 5) con-

taining characteristic branching pores 0.5 mm

diameter, alternating with clay, and sometimes

interbedded with silt and fine sand. The car-

bonates have unusual transitional or irregular

upper boundries: in some beds spherical zones

delineated by colour variations develop, which

pass upwards into discrete carbonate lumps

within the matrix of overlying unit. These are

thought to be diagenetic features associated

with lithification possibly resulting from inter-

mittent exposure. Other beds (Fig. 5) show

shrinkage cracks, into which the overlying

clay penetrates. Particles of carbonate are

included and flow lines occur, indicating

liquefaction resulting from thixotropic trans-

formation. The lack of rounding of the clasts

derived from cracking, and gradation to un-

cracked material, suggests sinking of carbonate

plates into underlying liquid clay. The cracking

may be a syneresis phenomena, which occurred

during or shortly after deposition of the over-

R. A. CALLEN & R. H. TEDFORD

NAMBA FORMATION

OUTCROP AT LAKE TARKAROOLOO

COONARBINE FORMATION

EURINILLA FORMATION

5YR5/6 +7

2,5YRS/6

5Y5/2-5/\

Stain 10YR6/2

5YR5/2 NI—N5

py Stain 1OYR8/6

ray

“4

fe)

—

Lat. 30° 07' 00"

Long. 140° 06° 34”

LAKE BED

74—956 Del, BT R. A. Callen S$. A. Dept. of Mines

Fig. 14. Namba Formation—outcrop _ reference

section.

lying clay. Occasionally the clasts have been

rounded, and incorporated in the overlying

unit: current or wave action has been effective

in some cases. Other beds show wispy car-

bonate and clay intermixed at the contact,

interpreted as flame structures which have

transformed by thixotropic changes, to flow

as a semi-liquid. Bioturbation is frequently

associated with these structures, and is com-

mon throughout (Fig. 6).

Unit 3 (49.7 m) is very similar to the lower

part of the Type Section (section 12, units

1-9), exhibiting similar cyclic deposition, in

which cross-stratified sands (Fig. 8) grade up

into tough black clays with pockets of medium

sand, often with polished grains (see descrip-

tion of unit 4, section 12). The black clays

are identical to those in section 13, Fig. 14.

Analyses showed the black colour does not

3 Ludbrook, N. H. (1972). Age and environment of deposition of a sample from Yalkalpo No. 1 Bore,

Lake Frome area, South Australia. §. Aust. Dept. Mines Rept. RB 72/207 unpub.

CAINOZOIC ROCK UNITS

result from anomalous concentration of car-

bonaceous. matter, sulphides or manganese.

Tron-nich montmorillonite or humic acid stain-

ing are alternative explanations. A bed of

dolomite or limestane nodules is often present

at the contact between the sand and black clay,

Lamination (Pig. 7) is generally not as pro-

minent #8 in the equivalent sirata in the Lype

section, and the sand beds are often burrowed.

The cross-bedded sand sequence of unit 4

(49.2 m tetal thickness) grades up into a

uniform olive clay with churned structure, The

sand bed is a promincat horizon west of Lake

Frome, and is being prospected for sedi-

mentary uranium of the geochemical cell type.

The dark sandy clays with skew planes are.

rather weakly developed in this unit.

The upper carbonate horizon, unit 5, is

23.7 m thick. has a much higher proportion of

clay than unit 2, and is intensely burrowed.

Sedimentary gypsum laminae are present.

The uppermost part of the section (unit 6)

in which the Namba and Willawortina Porma-

Uons intertangtte is mote conveniently des-

cribed when discussing relationships between

units,

The Namba Formation has been broadly

divided into two informal members (1 and 2)

of regional extent, on the basis of the presence

Or absence of |he tovgh black sandy clays with

skew planes. The lower member (e.g. units

1~4, section |1, Fig. 3) ts charactenzed by

these clays, and cyclicity is more prorninent.

It was later found that this subdivision closely

coincided with the change from smectite to

hte-kaolinite dominated clay mineral suites

(inset, Fig. 3), except in Yalkalpo | bore

(Fie. 3, séction 12), In this bore it is uncertain

whether the dominance of smectite throughout

the sequence fepresenis a local variation in

clay mineralology or whether the upper part

has Seen wrongly assigned to member 2

(which may have been eroded). The

mineralogy in Yalkalpo 1 bore is remarkably

uniform, smectite almost the anly component.

The higher proportion of silt is also unusual.

An interesting, varied vertehrate fauna is

found in the upper part of member 1 and the

base of member 2 of the Namba Formation

in various small saltpans southeast of Lake

Frome, in the vicinity of Eurinilla and Billeroo

Creeks, One of these Iccalities is at Lake

Pinpa (Scction 8, Fig. 15).

HILLAWORTINA FORMATION (Deriva-

timn—Willawortina Creek, passing south of

“Wertaloona" on the Balcanoona High Plains.

133

in the vicinity of the outerop reference sec-

lion),

The type section for this unit is Western

Nuclear’s sedimentary uranitm test hole WC2

(Fig. 3, section 1D and Appendix 1) cored

from & m to base, The hole was drilied on the

uplifted plains flanking the Flinders Ranges,

near Paralana, where a continuous sequence

of coarse poorly sorted sediments is encoun-

tered. A detailed division is not possible as a

result of moderate recovery and gradational

contacts. Three members are recognized, mem-

bers 1 and 2 (16.4 m and 17,06 m thick

respectively) have less mica and sand in the

mater than the overlying beds, and are less

oxidized. Member 2 has finer overall grain

size than member 3 but is comparatively

coarser than member one. Members 1 and 2

are equivalent to unit 6 of section 11.

Although bedding planes are very mdistinct,

transitions In grain size are often abrupt (Fig.

13), Secondary alteration with production of

red mottling is common throughout. Feldspars

are generally more abundant than in the

Namba Formation, Sandy beds have matrix-

supported framework with a high proportion

ef framework compared with the Namba

Formation,

The Formation crops out along crecks in-

cised into the high level plains flanking the

Flinders Ranges, along ihe southern shore of

Lake Frome, and along the Siccus-Pasmiore

River. The section (Fig. 3 section |, Appendix

lL) in a low range of hills, 3.7 km on 22°T,

north of Prism Hill and south of “Wertaloona"

{Air photo reference: § Aust. Dept. Lands

Svy, 803, Baicenoona Run 7, photo 0014), is

an important supplementary section, aS it is

the only outcrop in which the contact with the

Namba Formation can be observed. The

sequence is 140 m thick and dips 30-50° cast,

in accord with the remainder of the Cainazole

section, The whole rests with angular uncen-

formity on Middle Cambrian cocks. Exposure

is moderate to poor, necessilaling reconstruc:

tion from several scattered outcrops, particu-

jarly through the Namba Formation. This

sequence Was fitst mapped by Leeson (1967)*

who referred the conglomerate to the Telford

Gravel (Firman 1963, 1964, 1966b, 1967a,

1970) and the underlying clays to the Avon-

dale Clay (Firman 1967a}, Subsequently Callen

tin Coats 1973) remapped the area during

1970-1 for the COPLEY 1:250.000, zenlogical

miap sheet, and the sequence was assigned to

an undifferentiated Teriary-Quaternary unit.

134

Elsewhere on the eastern portion of COPLEY,

green clay, now known to belong to the same

sequence, was called) Avondale Clay,

In Sectyon 1 (Fig. 3) the base of the Willa-

wortina Formation is placed at the base of the

Jowest conglomerate, Beds below this unit

include poorly sorted sandy clays, but with

interbedded mictitic white dolomite, fine yel-

low-green sand, and pale grey and olive clay,

closely resembling the Namba Formation.

Below these beds, resting with angular uncon-

formity on the gently folded Middle Cambrian

red beds, is coarse sand with polished pebbles

and ?ferricrete clasts resembling the Eyre

Formation.

Another section regarded as eguivalent to

the Willawortina Formation, but of ayeral!

finer grainsize, is exhibited by unit 6 of Wonl-

tana No, 1 bore (Fig. 3, scction 11). It shows a

prominent. alternalion of sand and clay in fin-

ing upwards sequences, cach separated by sharp

egntacts, Sorting is uniformly very poor, and

matted green and brown colours common.

Secondary carbonate nodules are present, and

also beds of lacustrine dolomite. Toward the

top of the section the fining upwards sequences

become poorly defined, The top is capped by a

thin dolomiie bed, overlain by cobble con-

glomerate and sandy clay silt, representing the

Evrinilla Formation and “unnamed conglo-

merate’ {probably equivalent to the Millyera

Formation )-

Upsiream from section Z along Balcanoona

Creck, excellent exposures (e.g. Fig. 12} of

the upper part of the sequence seen in section

11 are displayed in cliffs. One of these exhibits

a hiatus—limestone and conglomerate In. the

lower part have been faulted before deposition

of the overyling silts, Subsurface (below soil)

karst structure ts present.

RELATIONSHIPS BETWEEN

FORMATIONS

The ature of the contact between the

Namba and Eyre Formations, and difficulties

associated with differentiation when both units

are sandy, hive been discussed by Woptner

ef af (1974), The disconformable relationship

is demonstrated palynologically by W. K-

Harris (pers. comm. 1974, sce section on

AGE, this paper).

The intenongurng relationship between the

Willawortina and Namba Formations is ilus-

trated by Fig. 3 (imsct). a section across the

Panlana High Plain, on which Wooltana 1

bore has been superimposed, A similar section

R. A. CALLEN & R. WH. TEDFORD

showing the same [catures can be drawn

across the Balcanoona High Plain through

WTS, WTS and WT bores (Mines Admini-

stration Pry Ltd) and Wooltana I bore. The

decrease in course clastic: proceeding east

from the Flinders Ranges t% demonstrated,

The lower boundary of the Willawortina For-

mation has been drawn at the base of the

characteristic mottled, immature. poorly sorted

sediments. Note the varying electric log re-

sponse to similar lithological differenecs be-

tween bores, which results from differing

drilling mud properties and sensitivity, and in

the case of WC2 bore, different instrumenta-

tion. Holes F22-20 and E20-13 however, are

not affected by these variables and are directly

comparable,

In Wooltana | bore (section 11 Fig. 3)

intertonguing with the Namba Formation is

exhibited by unit 6. The typical Namba For-

mation lithology of sharply differentiated

relatively better sorted clay and silt beds

grades to the extremely to very poorly sorted

coarse grained Willawortina. Forination, The

two units alternate Lo some extent. Essentially

there is a gtadual upward increase in the

coarser grained fraction, though an isolated

pebbly bed appears fow jn the sequence. Clays

are rich in illite (muscovite) and feldspar Fs

abundant, compared with the bulk of the

Namba Formation where these minerals are

minor components and smectite the dominant

clay mineral.

Dnit 6 of section 11 is therefore interpreted

as the equivalent of the lower part of the

Willawortina Formation in section 10, a rela-

tionship suggested by the correlation Jines

drawn in. Fig. 2 of Callen (1976). The criteria

chosen here to identify the base of the Willa-

wortina Formation are those readily mappable:

the base of the consistently coarse-grained

poorly sorted sediments, Thus unit 6 as shown

on big, 3 is regarded as mainly Willawortina

Formation, though il contains tongues of lacus-

trine dolomite like those in the Namba Forma-

tion. The contact is readily recognizable from

petrophysical Jogs (Callen 1976 Fig. 2} and

can partly be explained hy the degree of

secondary altcration (carbonate nodules, iron

oxide mottling) striatigraphically associated

with the Willawortina Formation. These

secondary cffcets alternated with deposition,

and are an integral part of the unit,

Support for the inlertanguing relationship

between Namba and Willawartins Formation

CAINOZOIC ROCK UNITS

is aso derived from clay mineral analyses

(Callen 1976) Results are shown diagram.

matically on the inset of Fig. 3 demonstratlig

the abrupt change from rocks dominated hy

smectite and randomly interstratified clay, to

illite (largely well crystallized muscovite),

randomly interstratified clay and kaolinite.

This change corresponds to the position of the

tlunite horizon within the Namba Formation,

and is widespread throughout the basin, having

been focated in 14 bores and in outcrop. The

change was probably initiated by uplift of the

Flinders Ranges, probably with climatic varia-

tton frorn high to low rainfall as. indicated by

clay mincrology and colour change (see later).

It is therefore regarded as an approximate lime

marker, and is coincident with the boundary

between members 1 and 2 of the Namba

Formation, and with the base of the Willa-

wortina Formation in its type section. The

change corresponds with the base of the Willa-

wortioa Formation identified in WC2 and

Woollana 1 bore.

Alunite is recorded near the top of member

1 of the Namba Formation, forming a serics

of nodular horizons associated with sharp

bedding planes. The nodules ramify through

the clay and resemble calcareous hardpans of

soils in their manner of development. The

horizons ate widely developed in the Paralana

High Plains area, but are also fotind in the

¢astern part of the basin in C1S bore. Here,

they are overlain by a relatively thicker

sequence of member 2 than in the high plains.

The horizons arc regarded as soils, associated

With a well developed hiatus or disconformity

formed during uplift of member 1. This em-

phatizes the time significunce of the clay

mineral change recorded earlier.

Silcrete has been identified by one author

(R.A.C.) in the interval 72-94 m from cut.

lings of bore LBIZ, drilled by Mines Adminis-

tration Pry Ltd, It is developed on ctay, and

ovetlain by greenish-+red mottled sandy cal-

careous clay resembling the Willawortina

Fotrsation. A number of closely spaced bores

beiween “Murnpeowie” and Reedy Springs,

drilled by Peechiney Exploration (Australia)

Pry Lid (Mannont & Barral 1972)6 suggests a

similar relationship. The silcrete vartes from

the red und grey mottled chalcedonic and

opaline “‘puddingstene’ to the grey rmero-

crystalline quartz “groy billy” type, according

195

to whether clay or sand is siliciied. This is

displayed by Mannoni & Barral in their cnoss-

section, and can he observed in outcrop, The

same siletete horizon fennms a cup to the

dipping Eyre Formation ac Reedy Springs

(Woptier et al, 1974 Fig. 2), The silerete is

thought to represent a soil horzon, and there-

fore marks 2 disconformily (Callen 1976)2,

Thus there is eVidence supporting a discon-

Tormity between the Namba Formation and

rocks resembling the Willawortina Formation

in this area, Althougl the silcrete has fot been

identified in the high plains regions, it ts

apparent that the Willawortina Pormation, as

defined, may contain some younger material.

The brown silerete and ferruginous material

developed on sandy facies of the Namba For-

mation exposed at Lake Tarkarooloo, and

around other saltpans east of Lake Frome,

are thought to be cquivalent to (hat just de-

scribed, Cementation certainly occurred prior

to deposition of the Millvera Formahon, as

indicated by abundant silerete nodules and

ferruginized Namba Formation clasts in the

base of the channel facies in Lake Tarkaroolon.

AGE

The flora of member 1 of section 11 (Fig.

3) indicates an carly to. middle Miocene age

fur the base of the Namba Formation {Bates-

fordian-Balcombian—pers. camm.- W. K.

Harris 1974). Harris states the fiara is stmilar

to that of the Munno Para Clay of Lindsay &

Shepherd (1966), and Lindsay (1969, p. 38)

in the Adelaide Plains Sub-Basin, An assem-

blage of the same age was found in Mines

Administration Pty Ltd LCIA bore (for litho-

logics] deseniption see Wopfner ef ul, 1974)

to the oorth of section }1. and also in Luke

Evie 20 bore (Johns & Ludbrook 1963) ijn the

Etadunna Formation,

The age af the Waillawartina Formation,

accepting a conformable relationship with the

Namba Formation, is therefore medial Mio-

cene or younger. Its upper ave limit. as for

the Namba Formation, is deduced from rela-

tiggship ta the Millyera Formation, and

Eurinilla Formation (Pt Il) indicating a mm-

mim age in excess of 40000 years B.P.. pos-

sibly pre-Pliacenc.

4 Maunnoni, N.. & Barrul. J. M. (1972).—Murnpeowile Project 5.M,L. 373 (South Australia) drilling pro-

gram report R/72-21-U, S$, Aws!, Depi, Mines envelope 1327, Unpub.

R. A. CALLEN & R. H. TEDFORD

136

sau jo idaq -y 5

“AB|A WOR \sOuaddn yy)

PIWWICSSE SUOZIOY |X) HOS AIUN}Y

“DRMDIS. BY 2WADLY LO YMG MO||BA

Snduidtiiay AQ padded sauiialuog

“SoMUEY SJEPLI|Y 40 (NBII Wa|sim

AuQ\E poynejan sadvanbas Suiddip

ge Ajuniuiads $837) ‘aWOL4 aYe7

{0 JS¥HUINOS PUB YBa “SHIEK

Pur Saye| Alp Jo saapA Auge

*squN 70a ajeuoqed

AURPUGIBS Ju ajasvdAd ‘3p

Aq padded Ua}jo 344)/9

SUSE MO}RYS PUL JOYUELY peautsepim ly

‘BRIN DeS|

Uo Ives Yyoyed HORA 'Wydap ype

jo SyStuy galssepy “uwtipas Snolod

Sus 9p9!Te) (ayeMpUNVIT

{nq PEW Sains 9 yyosid

“#yaays jeueziny Jo speaipuyyés

flugy un gadajanap areungJeo

J ABSA) PATO|RAAD {IAM

SOS dILVIOOSS¥

"au AHR]

= TOR PIE SuBALy

Mowsey Smyats Zuoje syijp "syed

BuG|e $49 TUFUPYAO JO (PII,

SP ya sdOsy WAVE smethjernz Ww

{ymol8 Siladdns \/ ay ‘sazuey

NOISS4WdX3

JIHddOW0I9

“

AYLAW049

SHLLYadOUd LINN HOO AO AUVWWNS ‘T ATEVL

tajind pur savods yuejd wos

(NYIBHODTWa-N¥iddOssalva)

(NOILYWYOI VaWWN

>} dysuoyeres WH) SNFIDOLSII Id ATW

Agwsscd ANADONdINAIOIN = 3LV7

ANAQOIW TVIGsW

yaoo yy

ed ed

"ARANRULINSHD “ARADEIO)SAY

'BRQUILI2I) “seepadpmangz

sAMesepad ‘wWinpAer

‘sndepayiuny ~S3iD0S |UE|d

“ROR

‘we /PLONGLEY "BYE IA OU,

‘sany ‘BYIpNAN]

RUUD NSOaa, “}OUdIG

Ops snfudctameyoy ‘Spi n0U

WIS ple PIUDOD salFIRI|

“(SmMUMIAQ) BpovE,|SO,

“YPAL ely BUnj—sayeaqaye,

“$peq Jamo) Ui sauids \ysl4

“SBIMpIOI] 12 \nBadst YyyM

Aeja Y3/Q Eno} ‘sayWojap

'sples pue syis auy

PUR SHES Laas haIT Arey

quaunuioud

WiniiSodap dXyoAQ ‘Sauunis

tie syoeta (snoonbeans,)

BBRYLYS Biey)

‘YOWUAd Loleqanaig

“YOUUWOT. SAINTS

May Hain “U

(evoneulznyesju) ‘padi

Gayo ‘sa}e

H|NA81I

li WOW Yulppsg-ssors.

9/205 WNIPAU UT [jews Als A,

‘PUES AU, AIDA. PUB

Ys Uy Aeradsa. ‘youiiion

LOBuNMe) jB}Uoz 104

Li99S FUE CS OILEADS Ele e-ENOES

, “WL GAS Ate “2 HL CHAD

BNON 9 C7 A/G ZAO7 1 “aUYysoLUy| SalE ley

BIBY ‘SSAUETUCW UIA PAU|} Se1od Mujyoueg Ag

fajenaved ae PUE ‘O/N:ONd}5 Is/NPOW ere) Ua}io

SAWL/ON “Andazeutgiks ay Ae SABI pi

“Aald 10 upala eyed

JO svinBue AjjesausA spues “aIGo Wau ‘spaq

SYWOIOP AU!) PUL ayO}Op wiy) ‘BK puE. ps

‘SpuRs payos AUOOd Winipaw oy uy Bulevsayy

‘18 [2G BSa—¥2

NOILVWHO4 VEWVN

“SRE, WEY

S1ISS04

NOWLWI3WY09 Od

VIHALIED

salae) dau) ‘sed

43M{ LY SapAD seman

SUIULY POL] Wf Spag.

‘sparp ‘$5022 apbas Uppal

paUYaD |18N\

pee vem apg

SIUMLONULS

AXVLNGWIGS

BG SALG 2 UR EMAL TA

y8ea Ja jUMp Epa pubs 'saTUeY S9Pl|4 Wea

i) 2d) Dy sagged jo Sasuay

SHOP Kapinan ‘sajqqad pasyye9s

UM FP WIPED alum ysivaBa alee yO $18AYs PLE

0 FA uoyayays

AU RUEW Tiddfarua saynpan ayenened

YsWwea!d squall aim Aeya yes AQnad

Gp AlaD|NOD PANE AWDOd Alem OY AyaUIa eT

ADOTOHLIT

NOILVAOS

VNILYOMVTIIM

CAINOZOIC ROCK UNITS

REGIONAL CORRELATION

Relationships with other units used on adja-

cent South Australian Department of Mines

Geological Atlas Series map sheets (COPLEY,

PARACHILNA) and in other basins, are

shown io Table 2,

Equivalence between the Etadunna and

Namba Formations is demonstrated by litho-

logical similarity, similar flora, and occurrence

of species of fossil marsupials previously

known only from the Etadunna Formation,

Both contain the unusual dolomite-palygorskite

mineral assemblage.

A sequence penetrated during drilling opera-

lions by Carpentaria Exploration Pty Ltd

immediately west of the Ediacara Fault { Binks

1972) is Very similar to that encountered in

Wooltana No. 1 bore (section 11, Fig, 3) in

the Lake Frome ares. The section in Binks’

Fig. 3 has been interpreted by one of us

{R.A.C_) thus: 0.0 to 94.8 m 7Willawortina

Formation equivalent, 94,8 to 121,0 m—un-

named beds, 121.0 to 233.2 m—Etadunna

Formation equivalent, 233,2 to 298.7 m—Eyre

Formation, The sequences in the intermontane

Walloway and Willochra Basins {Howschin

1909, 1913; O'Driscoll 1956) are more diffi.

cult to compare lithologically, but palynology

(Hartrs 1970)7 from 30 m in Willochra No, 2

bore suggests most of the sequence is equiva-

lent to the Eyre Formation,

On the northwestern side of the Flinders

Ranges is the Avondale Clay (Firman 1967a)

of similar lithology and mineralogy to the

green clays of the Namba Formation and

Willawortina Formation, patticularly where

they intertongue (unit 6, section 11, Fig. 3).

The type section is affected by secondary iron

oxide mottling, and the "clay" is actually a

clayey fine sand, with angular shiny grains,

The relationship between Avondale Clay and

Ejadunna Formation is unknown at the type

area: the base js not exposed, and the unit

is unconformably overlain by the Telford

Gravel and "Conglomerate at Lyndhurst”

{Firman 1969), The “Conglomerate ai Lynd-

hurst’ resembles conglomerates in the Wills-

wortina Formation (Fig, 12 this paper).

Kaolinite ig the dominant clay in the Avondale

Clay type section, and is abundant in the uppet

part of the Namba Formation, and the Willa-

wortina Formation,

L37

A section of Yetila Creek in the Mooloo-

watana area of the northern Flinders Ranges

owas tesccibed and figured by Firman (197i,

Fig. 12)8 as Avondale Clay, Upstream from

this site, a lower part of the section is exposed,

connected by continuous outcrop, This exhibits

micritic carbonale nodules, underlain by silty

olive grey clays similar to the upper part of

the section, The clay is capped by a well-

developed hard white fine grained carbonate

soil horizon, Comparable to that developed on

the Willawortina Formation at Balcanoona

Creek, It. is overlain by the Eurinilla Forma-

tion. The lithology is identical to the transition

beds between the Namba and Willawortina

Formations (section 11, unit 4).

The Avondale Clay is regarded by

as much younger than the Etadunna

tion, henee younger than the Namba Forma-

tion, However, the comments ahove suggest

it could either be part of the Etadunna Forma-

tion, equivalent to the lower part of the

Willawortina Formation or upper Namba

Formation,

The lower part of the Telford Gravel (Fir-

man 1967a) may be equivalent to at least part

of the Willawortina Formation.

Firman

Forma-

ENVIRONMENT

Consideration as to whether the Namba

Formation sediments are marine, marginal, or

non-marite was a prime objective. The most

conclusive indicators of marine influence are

marine fossils and glauconite, hence samples

were investigated for foraminifera, and any

green pellets or clays were studied by x-ray

diffraction, A variety of lithological types from

subsurface and oulcrop were examined by

J. M. Lindsay who found no foraminifera,

Non-marine gastropods and pelecypads. (pers.

comm, N, H. Ludbrook 1973-+) are present,

aa are non-marine ostracodes (pers. comm. K,

McKenzie) and the fresh water algae Pedia-

strum, (pers, comm, W. K. Harris) and

charophytes. Al] green clays proved to be

montmorillonite, and green pellets Tound in

The eastern areas were dolomite, associated

with non-marin¢ pelecypods.

Other evidence for non-marine origin 1s

derived from the terrestrial vertebrate remains

(e.g, Fig. 10). Several skeletons were found

in a partly articulated state, Delicate bones are

2 Harris, W. K. (1970). —Palynology of Lower Tertiary sediments. South Australia, M.Sc, thesis, Uni-

versity of Adelaide. {unpublished}.

§ Firman, J, B. (1971).—Regional stratigraphy of surficial deposits in the Great Artesian Basi and

Frome Embayment in South Australia, §. Aust. Dept, Mines Rept, RB 71/16.

R. A. CALLEN & R. H. TEDFORD

138

saulw jo ydaq ‘y's

NOILYWYOd JYA

N311V9' ¥"U

NO|LVWHOd FaA4

NOILYWHOd

VANNAOVLa

IDVIANS CINGIOD

AO 414dD1S

NOILVWaOd VaVENOO

ANOIS3AWIT YOYIITV

J1qadD TIS

o@r—rZ

INID0I1Vd ONY

3N3903 1VIGIW

INJJOIN ATYVI

©} 3N3909110

NOILYWeOd J8AI

ayaiajis Alpiq AaiB

IDUWN|OS SAISSOW;

INIDJOIN = AVI

} IIIA

NOILVYWHOJ VaWYN

ALJaD Naa

NOU YWaOd TAIT ydIM

QNY¥S NGYIOMNdWYW

2143 9427 1961

"}2 12 NOLMILS

SAD|>

4nyupudy yo

ajDJaWo)Bu0y, pup

AV1ID JIVINOAY

AV1D JIVONOAY

{ua}DAInba

é 343531143} UO ‘ajasd|/s

Al|iq Ao18 puo auojsBulppng é

INIIOLSITd

Aa]

} INJIOIW

NOILWWaOd

YNILIOMVTTMM

DIPDAsMY YNoS

usaysDayjsou puo

sasag Nypayazyg

‘'yZ6L ‘YANIAOM

Om 1,

INOLSIWI)

¥VINNNAONNE {@2Dj1NS OpPUooiDy fo)

*sajDsa wo) Buoy

uvoz1ioy snour6nisey

saays dow

000 OS2‘L AIIdOD

8 1? SIVOD

saBuoy

S1@PUI]Y WIASSAA “Nh

“NYWald

ajeidiiiay

jaded sy)

QdOs3dal F NIV

SNISV@ YSHLO GNV SLASHS dVW LNZDVrav - SLN3TVAINOS G3LS4a99NS

SLINN 43010 - LHVH) NOILVTEYIO

VWauV

3WOYd IVT

C 718Vl

CAINOZOIC ROCK UNITS

well-preserved, and abrasion due to transporta-

tion i currents virtually absent. The sediments

in which they occur are fine sand, clay, and

dolomitic clay. A nearshore marine environ-

ment therefore seems untenable, though a

lagoonal or upper estuarine enyirenment is

possible. A mon-marine environment is pre-

ferred, though presence of Cetacean remains

(a platanistie dolphin} indicates a link to the

sea at some stage.

More specifically, environments are (i)

Micritic delomitic carbonates with irregular

ooliths, suggesting low energy shallow lake or

shoreline conditions. ¢ij) Black, laminated

fossiliferous clay of Wooltana No, I, unit 1,

suggesting a well developed Jake: the fine

laminae resernmbJe varves, but have been dis-

rupted by diagenesis and bioturbation. (iii)

Sedimentary structure types, abundance of

fines, and very poor to poor sorting support a

low energy environment for the whole unit,

This may explain the apparent lack of well-

developed beach sands, which would be poorly

developed and poorly sored along a low

energy shoreline,

The environments represented by the cyclic

sequence described earlier are in ascending

order: (a) channels: smal] to medium seale

cross-bedded fine to medium sand (Fig. 8),

(b) flood plain, estuarine or lacustrine: finely

laminated silt, often burrowed and with very

small to small seale cross-bedding (Fig, 7),

and olive clay, (¢)} lacustrine: patchy car-

bonate, oolitic dolomite (Fig. 5) and clay

(Fig. 6). (d) swamp or mud flat with ovca-

sional channels: hard, black, niottled clays

with irregular fractures and sand patches

(Fig. 4), interpreted as vertisals. The cyclic

sequences are of 1-20 m thickness, averaging

9m, well-developed in most parts of the basin,

except the northwest where uniform clay sec-

tlons dominate, The cyclicity suggests a depasi-

tional process resulling in a particular sequence

of facres, but with inherent tstability, Some

examples applicable to the Namba Formation

are (1) a delta building into a shallow lake or

estuary (necessarily shallow because the cyclic

sequences are thin), (2) repetitive transgres-

sion and regression of a shallow lake shore

In tesponse to fluctuations in water level ¢3)

repetitive avulsion of a meandering stream,

(4) bars associated with development and

abandnoment af portions of mver channel,

The abundant bioturbation, and its occur.

Tence in medium-scale cross-bedded coarse

Channel sands, aud basal parts uf laminated

139

silt beds, is inconsistent with river channel

origin of these facies, These salids more likely

represent offshore lacustrine bars. Lenses of

the coarser sand facies at the base of, or with-

m, the tough dark prey clays are also difficult

to explain in terms of a river and flood plain

relationship: channels cutting across a tidal or

deltaic mud-flat are more acceprable. Subse-

quent intensive bioturbation or rheotropic flow

has partly destroyed bedding, distributing the

sand in irregular patches. Im the estuarine

case, the absence of any evidence for a marine

influence, parlicularly in the microfauna,

indicates deposition in the uppermost Teaches

of the estuary, In sequences where coarser

channel sands are interbedded with non-hur

sowed Jaminated silts, the river-channel and

food-plain relationship is still applicable.

Fluviatile deposits are abundant, and fossils

(e.g. Fiy, 10) of aquatic vertebrates (Dipuai,

Teleostei, Chelonia, Cetacea} suggests a per-

manent water supply, and fossil plants (Notho-

fagus and Podecarpus) indicate bigh rainfall.

The distribution of lenses of channel sands

within an essentially clayey sequence is typical

of meandering rivers, Al€hough only 42 current

directions were measured (mostly in the Lake

Tarkaroeloo area) resulis sug#est a southerly

component of transport direction for the upper

pant ef the Namba Formation, in marked con-

trast with the north-casterly direction of over-

lying units.

Dense vegetation is suggested by the paly-

nology of the basal unit 1: the modem des-

cendants of the species represented typify

rainforests. Abundant grass pollen are evidence

that grassland occupied extensive areas, Thiss

rainforest was not coniinuows in the carly

lacustrine phase of deposition. The relative

ahutdance of arborcal marsupials in the upper

part of the Namba Formation indicates the

presence of gallery forests along the water-

courses.

Apparently at variance is the smoctite—

dolomite—palygorskile association, frequently

recorded from arid soils. playa lakes and warm

hypersaline waters (e.g McLean ef al, 1972,

Bentor ef al. 1963. Meester 1971, Singer ef al,

1972), At present dolomites and high-Mg

calcite are forming in hot arid or semi-arid

hypersaline lagoons (e.g. Von der Barch 1965,

Von der Borch ef al, 1975, Fricdman et al,

1973) though some magnesiumerich sediments

are found in fatitudes as high as 48°N (Muller

er a. 1972).

144

Millot (1%64) indicates the montmorillonite

—palyvorskite—sepivlite association is the

result of offshore lacustrine or marine chemical

deposition, This toek place adjacent toa laleri-

tized land mass of low relief and dense vege-

tational cover, in # subtropical or tropical

climate, Sepiolite is absent an the Lake Frome

area, but this may be an affect of degree rather

than basic difference, The hypothesis as applied

ta the Namba Formation overcomes the diffi-

culty of evoking evaporative conditions in a

high rainfall climate.

Millot’s hypothesis has been applied in a

similar manner to the Cainozoic rocks of the

Jordon Valley (Wiersma 1970). These sedi-

ments contain a remarkably similar sequence

of clays to the Namba Formaticn, Particularly

televant are Wiersma's comments regarding

ibe ongin of palygurskite {p. 88). He con-

cluded that intensive weathering on the hinter-

land under warm humid conditions was

necessary for liberation of the clements essen-

tial to the genesis of palygorskite and its

associated sediments, and that evaporation in

the sedimentation basin should be such annu-

ally as to provide the necessary concentration

of chemical elements. He deduced that

evaporation must prevail over precipitation

and fuviatile and/or marine supply of water

to the basin. In many places in the present

tropics evaporation can exceed annual pre-

cipitation, with resultant formation of evapor-

ites in favourable locations, Palyvorskite was

of detrital origin in the Jate Tertiary and

Quaternary of the Jordon Valley, having been

derived from Cretaceous and early Tertiary

recks im which it originated by chemical sedi-

mentation. In the Lake Frome area oo pre-

existing rocks rich in palygorskite were present:

rather kaolintte, smectite and illite are ahun-

dant. Palygorskite can be formed in soils

(Singer & Norrish 1974) hue only in relatively

low proportion, thus it mmnst have originated

within the depositionary basin during sedi-

mentation.

(tis notable that Mellot’s 141964) ideas as

applied to deposition of Namba Fartnation

stdiments require on equivalent Miocene

fateritization on adjacent land masses. In this

context Wopfner’s (1974) conclusions regard-

ing an Oligocene-Miocene “ferralitization” are

of interest, Although the evidence he gives for

age of the Doonbara Formation is inconclu-

RK A. CALLEN & R. H, TEDFORD

sive, some ddditional observation are made

here, Firstly clasts identified with the Doonbara

Formation (by R.H,1,) are found in the Wipa-

jiti Formation (Stirton e ai. 1967) of Miocene

age, Secondly the ferruginization in Lake Eyre

Bore 20, doubdtfully equivalent to the Doonbara

Formation, 1 recorded by Callen (Wopfiner

er al. 1974, Fig. 17) within the lower part of

the Miacene Etadunna Formation. Others have

also recorded an older Testiary ferricrete ( Fir-

man L967b), Therefore lateritization {or at

least, ferruginization) could have been pro-

ceeding in uplands adjacent to the basins in

which the Etadunna Formation and Namba

Formation sediments were being depasited.

The main carbonate hotizons occur a few

metres above unit 1 of sectlon 11, with its

rainforest flora, and above the vertebrate zone

with its indications of seasonal climate with

abundant water supply. The presence of these.

carbonates can be explained in terms of pro-

tracted urid phases superimposed on a sub-

tropical or warm temperate climate.

In addition the presence of detrita) feldspars

must be explained, particularly in view of the

abundance of plagioclase and wssociation with

smectite." The possibility of addition of vol-

canic material From eastern Australian must be

considered, the Miocene being a period of

Maximum vulcanism (Sutherland er af. 1973).

However, the percentage of feldspar is not

large. Preliminary studies of feldspars im the

Namba and Willawortina Formations suggest

Telative proportions of feldspar types and

compositions of plagioclases are similar to an

unmodified contribution from nearby Precam-

trian crystalline basement rocks. On present

evidence there is apparently no change in rela-

jive abundance and type of feldspars in the

illite-Kaolinite rich zones of the Tertiary, in

comparison with the smectite zones. This sug-

ests abundance is not tied to smectite occur-

rence, as would he expected if these minerals

originated from volcanic ash falls, The pre-

sence of feldspar presents a problem consider-

ing the evidence for a humid climate. Possibly

seasonal aridity and nearby source permitted

preservation. In addition Todd (1968) has

shown plagioclase is more stable than ortho-

clase under conditions of restricted leuching,

in a tropical climate, Thus smectite (mont-

morillonite) is unlikely to have originated

from volcanic: ast falls,

The mineral proup smectite, but R. N. Brown recovnized dioctshedral montmiorillonite in several in-

stances.

CAINOZOIC ROCK UNITS 14h

In the final analysis, the Namba Formation

was thought to be deposited in o warm tem-

perste to subtropical climate. The landscape

ad a savannah aspect, With gallery forests

around permanent rivers and lakes, Periods of

aridily occurred.

High average temperature, invoked to ex-

plain the mineralogy of the Namba Fornatinn,

is in accordance with marine paleotemperature

measurements mm southern Australia (Gill

1968). and New Zealand (Devereux 1968,

Jenkins 1968) of 18-22°C for the Miocene.

Considered in the light of continental drift

data, which suggest Avstralia was closer to

Antartica (though drifting rapidy northward:

Wellman e? ul, 1969), and data which indicate

the cooling of Antarctica way underway

(Hayes ef al. 1973). the temperature can be

explained in terms of greatly expanded sub-

tropical climatic zones during early and middle

Miocene times.

Deposition of the Namba Formation in the

central part of the lake Frome area was fol-

lowed by widespread ferruginization and silict-

fication (opal, and quartz overgrowths) and

development of cryptocrystalline silica nodules.

particularly in the coarser Namba Formation

sands, These processes Were the result of wide-

spread groundwater movements, Formation af

dunhcnists ynd telated phenomena had a locus

in river valleys cut into the Namba Formation,

prior to deposition of the Millyera Formation.

No evidence for 4 major period of Oligo-

cene to early Miocene “ferralitization” sie-

gested by Wopfner (1974) was found in the

Lake Frome area, though there ts abundant

evidence for late Miocene to Pliocene ferruy

ginization and orthoquartzite silerete Forma-

tion. This does not necessarily negate

Wopfner's climatic evidence, since two periods

of fertuginizition are prohable (Firman 1 967h,

19714; Jessup & Norris 1971). The older

Tertiary ferruginization would presumably nat

be manifested in the Lake Frome area, where

chemical and detrital deposition were pro-

ceeding.

The coarse detritus in the Willawortina For.

mation has clasts derived from Cambrian and

Precambrian rocks in the Flinders Ranges,

When considered jn combination with poor

sorting and abundant feldspar content, vigorous

uplife of the Ranges is indicated. This wes

accompanied by movement on the Poontana

Fault. A similar conclusion has been drawn by

Binks (1972) from evidence on the western

side of the Ranges, Ironstone and silcreie peb-

bles from pre-Willawortina Formation (?pre-

Namba Formation) duricrust are present,

However, laterite clasis are got as abundant in

the overlying Willawortina Formation as ont

would expect in a sequence supposedly derived

from erosion of a laterized land mass, Pre-

sumably this is because the Flinders Ranges

were virtually non-extstent at the time of

deposition of the Namba Formation, presenting

only a smiull atea for laterization- Alternatively,

in keeping with the suggested warm-temperate

to sub-tropical climate, ferruginization may

not have developed an extensive laterife crust

Deposition in an alluvial fan environment is

suggested for the Willawortina Formation by

the presence of extremely poor sorting {Fig.

13), numerous channels (Fig. 12) with

medium scale cross-bedding, and laminated

calearceus sills (Fig. 12) with red-mottling

snd carbonate conerctions typical of flood

plain deposits. Firing upwards sequences are

typified in section 11, suggesting bar deposi-

tion, The deposils coarsen very rapidly close

to the Plinders Ranges. The extremely poor

sorting, coatse grain-size and matrix-supported

lexture in some beds may be the product of

mud-fows. The red mottling (‘marmorization’}

and carbonate soils are similar to those des-

cribed by Freytet (1971) in association with

alluvial deposits, and typically form in the

inactive parts of fans (Blhissenbach 1954, p,

185; Denny 1967, p. 105). These features

resemble modern fan deposits.

In sections | and 10 there is a tendency for

averall coarsening upwards, suggesting increas-

ihely rapid uplift of the Flinders Ranges. The

uplift deluged the former lakes and swamps

of the Namba Formation with detritus, re-

ducing their extent. Thin dolomite lenses in the

sequence (section 11, and Balcanoona Creek)

represent lacustrine or playa lake phases simi-

lar ta these of the Namba Formation, Petru:

logical investigation shows these contain 4

much higher proportion of sand (mutch of it

unstable mineral grains) than the Namba

Formation carbonates.

During deposition of the Willawartina For-

mation, oxidizing condijions became prevalent,

through accumulation of the sedimentary

column above the water table, This contrasts

with the sub-water table reducing eavironment

of deposition of the Namba Formation, Abun-

dant potash feldspar and plagioclase can be

attributed to rapid deposition and possibly

semi-arid climate. Presumably uplift of the

{42

Flinders Ranges Would haVe had a strong effect

on bocal climate, but this cannot be assessed

at present.

Following deposition of the Willawortina

Formation, ferricrete and calcrete formation

occlicred, particularly in marginal areas.

The absence of surficial cementation of

coarse sediment in the type section of the

Willawortina Formation and nearby outcrep,

contrasts with the ubiquitous cementation in

southern areas (sections 1, 11, Fig, 3). An

explanation in terms of a carbonate rich source

arca for groundwater or sediment, or abun-

dance of limestone clasts in southern ureas,

does not explain the widespread distribution

of surficial carbonate cementation in rocks of

various ages throughout the Fhnders Ranges.

Indeed, many fans in semi-arid areas through-

out the world are similarly cemented, Enough

calcium is produced by weathering, or de-

posited from wind-born dust, to provide suffi-

cient carbonate material for cementation

anywhere, Therefore in the case of the Willa-

wottina Formation adjacent to Mount Painter,

absence of carbonate is a local phenomenon,

the explanation of which is unknown.

Pt. [1—Younger Cainozvic Rock Units

Type and reference sections are shown in

Fig, 15, and described in detail in Appendix

2. Table 3 summarizes the lithological and

other properties of the rock units deale with in

the text,

LITHOLOGY

MILLYERA FORMATION — (Derivation:

Lake Millyera*, near the mouth of Billeroo

Creck, Millyera is Jocal aboriginal word for

water. Map reference: Siccus map sheet,

FROME),

The name is proposed for a sequence of

imerbedded greenish ostracode-bearing clays,

thin limestone of charophyte algal remains,

and fine sand, The sediments occur in Lake

Frome or io smal} lakes close to its margin.

The name is also applied to 9 coarse cross-

bedded or conglometralic sand, regarded as a

fitviatile equivalent of the clays, where these

contam interbedded charophyte limestones.

The type section at Lake Millyera (Pig. 13.

section 4) is located 69.2 km en 320°T, north-

west of Low Stony Hill (map reference Tele-

chie) on the Siccus map sheet (Air Photo ref. -

Dept, Lands Svy 31, Siecus Rum 1, Photo

4460),

The section consists of 4.3 m of laminated

green ostracode bearing clay with « thin bed

K. A, CALLEN & R. Ho TEDFORD

(40 cm) of laminated charophyte limestone

near the top, overlain by alternating clay and

fine sand, An abbreviated description is given

in Table 3,

The \ype section was not Jocated at the

thicker section 3, where there is intertonguing

with red beds, This avoids confusion which

might arise should the red beds, which have

affinities with certain fluviatile equivalents

(c.g. the Eursinilla Formation—see [ater), be

formalized as a distinct unit.

The contact with the Namba Formation

Was nol exposed, but can be observed in the

supplementary section located about 2 km

east (Fig. 15, seclion 5, 68.7 km on 327.5°T

northwest of Law Stony Hill—air photo refer-

ence S. Aust. Dept, of Lands Svy. 361, Siccus

Run |, Photo 4461},

In section 5, the Millyera Formation is

3.3 m thick, cropping out below a thick ex-

posure of Eurinilla Formation. Here thin

charophyte limestone in the Millyera Forma:

tion grades laterally into gypsum, often ripple

marked (Fig, 16), imtercalated in an essentially

sandy sequence. Sometimes botryoidal sirye-

tures are present on the surface of the gypsum

laycr, similar to those found on the floor of

Lake Frome. Scattered very coarse polished

sand grains are present on top of the gypsum,

where it is in contact with the overlying red

sand lens.

The red sand lens consists of u thin bed of

bright red-brown coarse silk with basal granule

layers impregnated with secondary gypsum,

closely resembling the Eurinilla Formation in

lithology. It grades by alternation to greenish

very fine sand with coarse sand, silt and clay

laminae {yellowish grey). The greenish sand

bed is fossiliferous, with numerous charophyte

oogonia and stem moulds, fish verebrate and

spines, and ‘Coxiella’,

The contact with the Namba Formation was

exposed by trenching. Orange and yellow sands

of the Millyera Formation contain reworked

and oxidized tough grey sandy clay clasts from

the underlying Namba Formation, The Namba

Formation is more indurated and darker

coloured.

The top of the sequence is marked by a

strongly developed soil, also observed else-

where affecting the Namba Formation. The

soil has a crumbly texture, with peds of

irregular shape about (,5-2 cm across. A well

developed black mungams is present, and red-

thsh-to yellowish-brown iron oxide patches are

developed in the clay. A similar horizon is

)

seul jo ‘deg yw $

‘RUN OW Ue. BL EO-pay

Spues sbiggs jeseq

eS dHuuNO UBNY EpURS Ul

1 L3}AMPUNOM 2p BALSSEY

‘SUDYSALAY

PUNY, “UbAISAS Yaa

Hj -C018) pum Frayer ¢

pajuaseute. MoU

Bue papdue deep

idngie ‘ofpa pyncue yoo

OW PVE FUDIY 2HE] U0 pag Hl)

way yd

eas eoqysyy woud)

“Susedap JeaL SIOMN|LAISIp OFeq DAES

AN3DOLSI9Td ATHV3-3NIDONW

*avaapiaN 4SI] “aARAI

FapPMRNSO

Wve). SXtingy Gary

in Jada Oy ari

‘ts

Spodtuyse8 (Ends

pue spqdun

elugbon sojeuty

YUM. PUES al

Suureg-ponensa ali

“AUO]S aI AINA]. Sappueeey

‘Suippaq ANE

UOELHUR) SUID B/RIS PLS

“Sples faulunys pappag-ssisa

aly SNH) SayeaBy

JD. WnSOKG We deysiyy

akptancg pues sydudu.

lS. aLyaLIASSE

aoa puE ye ul

UQUeUHUNE) | 2qUOLI LOA’ Ub

“spurs AKO) Ys\U83 olny OY Lupa

LVRS 22-940 G pues i

VALAT POL aUN DUBS aU PL/ BAS “ISAS

“2USMAUT ‘Agr “SULTS puna

UL pueY- fappaCAIN|

UIA AUES @UN( alm 17 ySIpAIY “OPUNOL |faM. OF

PINE Swe UUs “ypey WHIOAS. POxsPU-Naiddes

Wi, OY Apel Ae GLOOM TAA biLAZ)

Due PiaOs-ypayk aU UWA POpADIET “Coy

Je PEUIWUE CLG ASP HOS

ny vedy |PIGYALOI. ‘Agja- Paysiee

ia 90 é54—re

NOILVWYOS

VUYSATIIIN

“sdh) JB yEmWUNON

fo LINSAAR pue ayeuncquer

HOS 0} fey Um YM panuoWa?

Alpilos dyevauio|zL09 jesey

BUY PLE pod {Oo Sapeys ul

PaylOW Sayeyod wnsdse paiatey

10 BAISSEW YIM ais|y ‘aNloyes

SE) Ul palual

Aalan|s OY AYRAM, puws Jaap

PIRPINS UO SpHAyS Ad sojmuesd

$F [No Jayjean Ajluanting “segde

ALS Wt Sapnpost 981] 4yoe ‘Hay

vB Pajuasand aq ABW Splaupulj Aa

WINSUAR Dur epeungiea Jo

SUULDI| paOD|Ehay [fam AEE

‘Sayeun|

wnnday auaay S4e7 aos

“3°5 Booje spunoul pile spurys

“eeguien) Avjo jyesaid syeubxouder

SOUUEy) feseq Ayyinbesy

yd ay

pat

4e> anisseu 16 Youag 7

DRIED A TRIAD AG papules

ayo ped fama] “sasinoalyR pue

Viowssaudap punowe yedoys Mo] se

joo Buiddoy ‘Siejd TUiA)-miy Vey

—

uaLuipss 40

SUUUN|2D BINWIGY ‘mands AyobIq

JBAMUBLAY 'SAONLI\UOImIE “SERA

PiEY §2, Tayayjeam 'sproupu

ayeWog iP padajanap Ay4e:

SOS DALVIDOSS¥

“HUI ARRIETA S7TRp

yeaa Se sysauy Wi yno sdou

“Salma [J35 pile shied: jo adenspue)

SOpEUIGD ‘Suey FuMdj mc] Tey

NOISSIHdX3

OIHdHOWOI9

aReSqeNeh

“pues 91203. U)

ymioodsa ‘undo seyeresy)

SAAWINA) “ALOLy ye]

40 88pa Jee eyaxoy,,

SYOUEL ee) pur

SPMNONSED 10 s3A2] \ESED

OS sadkouzeyy

qeyad vopoydtoatss

“Os vopoqudie? Fi

Sqn) (ese sayeIgayIEY,

‘Sysau py SMALING pAPSU)

duyep Mig cuney

ANIDOLSIF Td SLYT

SMOKY SII "JadyS PURI PesiMapM sejnqe) UIE

plies Linsdég

“tied yeseq ul)

ayvodkd pus le a ajem

PUNE ‘suorLey es penthy

UT YSUMO|pAA. JEAND]

WMOIT-AIS\pIKOd Bd sidde)

“PappHG- S807 Ed 191K)

"UCI EINEE-SsULs

PEPUCELOLY Cues, jpeuepeape jy

SRY Ailes Gun JO

ANY Syne) "WT Sadiq

iad 150) wyeOD ted

Jn avy auiy10§ s005

“pues winsdAy

ib duippaq-ssoa

addj—ayjouiny ay

osysin) ped

leo ny

(ed cago) aley syeos

Walpaw vO pappsy St019

dpueyd 1a yfnoy

SPURS |ESEY “Slay sue MO]

AROS aH OPW) pachjarsp

Salvia (WIDE Z)

WOyeU re) pepe Oy

PauOJONda aH soipespreys

‘Spues Winsdé asaeiry “yseia Qug_SuulES sppuiBLLoy

DUE SjoUOGIET paruian Ayeoo] PhP sayqqad

SHEN YUM fluUkS astegg [LE SopefaloyAyuy eseE

BUIAS SeSMADL 2 wed. 1aMO] ApUES

“7 bleepYAS-GZ MEM sealily| "pAssou) Oe Ppl

JG “pagyld Pue arial yng, papureY AjayeapULY

at] APN SiNeAS ‘PayVOS fyudOd Ada, “saygqad

YJ Wap} pees pred Jamo) “Wen Jadan Wi Keys

PUR YAS LO Spagdsyur yay "puRS cy 7

{6'€'S)

NOILVWHOS

VaTINIaNSa

| SMAUN “Suyeiqayian HEH

(881) “ORDO RalRIRYT

S}OE UE 12

(ays MUR “SyRy jreus

suaper wo) § 1 NIDIW

~ANIIOLSIATd ATYVA

PequeyG wuy

(NOILYWYOS YOINIUNS

AY19W033 STISSOJ

SLINA

Joy pe vomOY

(ive jesnlAy “SHeUS puey

40 WOUARTE VNO|OD lmMOsG

1h AUS gyevapoW

“BUIP2Iq Yavjaanp- AYBaN,

NOILVTaHHO9 404

VidsLldd

WEMIOTINOY OPM JG Woy

‘Tpauiyaosjols Jyaues)

MONET ENS-S50N9

SIPAS, FAIR] ‘Rasta

IDE} \EqunUcy yRayy

SIMNLONYLS

AdVLNAWI03S

YAJINNOA

SEILYAdOUd LINN MOO FO AUVWWNS € AIAVL

nS "Baus Marezapow

UL ‘PUES Aake[a JO AIS 0) PrleR

ASOTOHLN

NOILWWHOS

ANISYVNOOS

(1 ‘Gl4)| LINN

144

developed on the Millyera Formation in Sec-

lion 7 of Fig. 15,

The distinction between the lacustrine Facies

of the Millyera Formation ani the superficially

simijae Namba Formation can be demon-

strated from Lake Millyeta and Lake Tar-

karoaloo. In Lake Millyera, the north shore is

formed by cliffs of Namba Formation clay and

dolomite showing no facies changes thrausheut

the length of the continuous outcrop. The

south shore has sporadic outcrops of Millyera

Formation, also unchanged along strike, traced

to the junction of Billeroo Creek and Lake

Tarkarooloo. These relationships suggest a dis-

canformity between the two units. The Millyera

Formation alsa occurs on the north shore of

Lake Millyera at its castern cd, but a covered

interval prevents direct establishment of a

‘clationship with the nearby Namba Forma-

lion, ‘The contact between the units can he

observed by trenching the base of the supple-

Tentary seclion 5, where the erusional contact

und weathered top of the Namba Formation

support 4 disconformity.

Similar faminated ostracode-beuring clay

and gypsum laminae are found bencath the

hase of the gypsum dunes constituling the is-

lands of Lake Frome. These grade down to

ostracode and charophyte-hearing indurated

elays, without sedimentary structures, beneath

the lake bed. Fine sand interheds are present.

These beds ate equated with the Millyera

Formation. but most sediment flooring Lake

Frome, though of similar lithology, is younger

and less consolidated.

Along the eastern edge of Luke Frome is a

series of eroded mound springs, exhumed hy

deflation of the modern lake Moor These are

built of saucev-shaped carbonate layers, partly

algal in origin, which intertongue with the

clays, These spring deposits are partly ccjualed

with the Millyera Formation,

In Lake Tarkaroolwo, reddish-brown silt and

conglomerate is inter-bedded with the charo-

phyte limestone. Proceeding south along this

linear Jake, the coarse facies eventually

dominates, the limestones being absent. The

Millyera Formution is no longer identifiable,

having graded inte an entirely fluviatile facies

of conglomerate 2-3 m thick, cemented with

massive white secondary carbonate (hence-

Forth referred fo as the ‘unnamed conglo-

menate’ cyuivaleat to the Milkyera Formation)-

Section 6 (Piss 15, 17) shows an intermediate

stage in this transition, Numerous well deve-

R. A. CALLEN & RH. TEDFORD

loped channels exhibiting 7 cross stratification

(Allen £963) are present, cxhumed on the

lake Noor, They often contaiii clasts of fossil

wood, Namba Formation dolomite, ferru-

ginized Namba Formation sand, und mitky

quartz at the base, demonstrating a discon-

formable cclationship with the Namba Forma-

tion. There is little facies change along the

length of Lake Tarkarooloo in the Namba

Formation, whereas the Millyery Formation

yavies considerably, though retaining — its

identity a5 a wnil.

Two charophyte Jimestone horizons were

developed in the southern part of Lake Tar-

karooloo, instead of one ss at Lake Millyera.

Since they must have once represented a

horizontal lake shoreline, and arc equivalent

to the horizon at Lake Millyera, structural

deformation (?faulting) is required to account

for their relatively higher position in the land-

scape. Barometric Icvelling, tied in with South

Australian Department of Lands bench marks,

established the height difference [see Sections

§ and 6, Fig. 15). Comparison of the heights

of equivalent Namba Formation carbonites,

between Lake Millyera and the Namba Formu-

tion reference section in Luke Tarkaroolon

(Fig, 14) also supports downfaulting of the

Take Millyera region.

The “unnamed conglomerate’ channel

equivalent of the Millyera Formation can be

traced throughoul the area southeast of Lake

Frome, where it is invariably overlain by the

Eurinilla Formation. The disconformity is

difficult tro detect away from low4ying areas

such as Lake Varkaroolon, where well-deve-

loped greenish carbonate nodules and cylin-

droids of a soih calcrete mark the contact, and

massive White groundwater catbonate cements

the conglomerates in the Milfyera and Burinulla

Formations. Elsewhere the conglomerate has a

weak earthy carbunate cement and interbedded

secondary gypsum Javers. interpreted as

proundwater phenomena rather than 4oils.

The contact with Eurinilla Formation appears

gradilional fez, units | and 2 of the Eurinilla

Fennation in section 8, Fig. 15), and it ts not

possible to establish a disconfarmity. An addi-

tional problem in the Millyera Formation is

the repetition of red sand facies resembling

those of the Eurioilla Formation. This suggests

it may be possible to have Iwo red facies super-

imposed, The contact between unit 2 pnd 3 in

section 8 may represent such a boundary (ie.

ihe lower part of this section may cnrrelute

with the Millyera Formation).

wy 20-—

wi 2? 3

= 4

- N 5 N 5

>. 15

Ee Baleanoona W E

> Creek .

Lad

w 10 ee aS, \ 8 ERA

* FORMATION z =\7,, MILLY \

= MILLYERA =

FORMATION J FORM. © ROE SS

5 = Lake Millyera

FAULT : Lat. 30°0232' NAMBA \

Long. 139°54'42 FORM. \

Lat. 31°02'59'

Long. 130°Sé 36

"\. EURINILLA

Let N 3

7 ae FORMATION

Lot. 30°53'30- and 30°56'0)"

4S Long. 140°1014° ond 140°1)'06"

BINE

cOONAR

FORMATION SSS SSS) |

]

ce)

| EURINILLA

FORMATION N\ SF POT Rea rae

YPF SECTION

Reet FORMATION

Le)

MILLYERA . DEX Lake Pinpa Lake Koorke

FORMATION = Sa = Se ee ee Se SS

Lot, 31°09 59" q Lol. 30°58 24°

Long. 140°|3'39" Long. 140°19'37

FORMATION

Lot. 31°0640"

Rou Long. 140°06'34

w NAMBA FORMATION

* SECTIONS BAROMETRICALLY LEVELLED

DATUM: DEPT. OF LANDS

BENCH MARKS 5062—5066

a REFER FIG, | FOR SECTION LOCATIONS

a REFER FIG. 20 FOR LEGEND

35 2 NUMBERED UNITS DESCRIBED IN TEXT

°o

30 F

ire

et)

L74—23 S.A. Department of Mines

R.A. Callen Geologist Orn. B.T.

Fig. 15 YOUNGER CAINOZOIC UNITS, LAKE FROME AREA

15 ee TYPE AND REFERENCE SECTIONS ‘

1100—46.76 ~G6808 O

CAINOZOIC ROCK UNITS

Current directions Were recorded from a

variety of cross-hedding types in widely scat-

tered localities in the basal channel Facies

(both sands and conglomerates), though with

a bias towards the Lake Tarkaroofoo area, hut

are sufficient (45 measurements) to record u

nocth to northeasterly transport direction, Con-

glomerates such as those of section 8, where

no disconformity between the Eurinilla and

Millyera. Formation was established, were not

included in this analysis.

EVRINILLA FORMATION — (Derivation,

Eunnilla Creek, Evrinilfa map sheet. CURNA-

MONA),

The type section (section 7, Fig, LS) is

located at Lake Moko* on the junction between

Billeroo and Eurinilta Creeks (Air Photo Ref.

—S. Aust. Dept. Lands Svy. 395, Coonarbine,

Run 3, Photo No, 9637). The locality is

45.8 km on 332°T trom Billeroo Watechole

(CURNAMONA—Billeroo Creek). and the

outcrop (Fig, 20} occurs in an amphitheatre,

formed by guilics draining into the north end

of the lake. The upper fine grained facies is

well developed. The complete section is de-

scribed in Appendix 2, Section 7 was con-

steucted from two outcrops 200 m apart,

correlated by following the beds along stnke.

This sequence consists of three units, separ-

ated by a weak carbonate soil horizon, The

lower unit, exposed in the southern outcrop.

consists of 1-8 metres of white well sorted

medium gramed channel sand. It i ¢ross-

bedded, and contains some vertebrate remains,

clusis of underlying units, and ¢harophyte

oogonia. The sands have features jimilar to

channel] sands elsewhere at or below the base

of the Eurinilla Formation. An interesting

feature is the presence of biotite, suggesting

nearby crystalline bascment outerup at the

time of deposition. Unit | is averlain by the

much more widespread and typical fed silly

and sandy facies of units 2 and 3, respectively

3.3 .and 5.5 m thick, These units are separated

hy a weak carbonate soil harizon, The lower

is paler than the upper, and shows tio evidence

of bedding. It grades to clay-silt at the hase.

The upper unit has diffuse large scale cross-

bedding, is sandy, and orange-coloured

throughout. It is capped by a well-developed

fossil soil horizon with nodules and cylindroids

of carbonate and some gypsum,

The sequence is overlain above the fossil

soll by a horizontally luminated sand, similar

to thar al the top of Unit 2. but with more

145

pronounced bimodality. Since the sail repre

sents a disconformity, these sands are excluded

{rom the definition, and placed in the Coonar-

bine Formation,

Another section 8 m thick (sectuon 5, Fig.

15) is situated on a steep blu? facing north,

on the southern side of Lake Millyera, close

to the Millyera Formation type section 4. A

detailed description is in Appendix 2, The

sands afe paler than those of the type, and

contain low angle cross-bedding al the base,

The overall two-fold division fof units 2 and

3) in the type section is retained. A diffuse

thin bedditig dips toward Lake Millyera. Con-

sidered with the geometry of the outcrop. the

sequence is interpreted as a small delta.

The upper part of this sequence has tubules

and cylindroids of soft white carbonate and

gypsum, several centimetres diameter, forming

several inter-related horizons representing =

fossil soil complex. The tubules weather out

as hard cylinders. The lower part of the

sequence is partly cemented with shects of

sofi white carbonate, and with pink carbonate

nodules. The base is solidly cemented with

gypsum. partly derived froni the dissolutiun of

gypsum lunetftes {represented by low angle

eTuss-bedding).

A third reference section of 3 m thick 1s

located al Luke Koorka* (Fig. 23). a small

chtypan on Eurinilla Creek, close to the

boundary between FROME and CURNA-

MONA on Evrinifle map sheet, The western

edge of the pan is formed by a cliff 6 m high,

where scction 9 (Fiz. 15) was measured

(Appendix 2). Here, the Eurinilla Formation

is represented by mottled very pale orange and

strong brown clayey silt withoue stratification,

capped hy a massive gypsum horizon with

0.3 cm rosettes of gypsum crystals developed

in red-brown silt and gypsum flour. It discon-

formably overlics the Namba Formation,

Burrowed horizons and gypsified roots are

locally common in the Eurinilla Formation,

though not represented i the sections de.

scribed here.

‘The carbonate zones ut the top of the

Eunnilla Formation in sections 5 and 7 are

regurded as a single widespread paleosel, They

differ from the soil developed on the overlying

Coonarbine Formation by having farger

patches of carbonate segregation. often in

several horizons, [requently weathering out as

solid sheets or lumps, In the Arboola Claypan*

large soft calcureous “biscuits” are developed.

In which the onginal lamination of the

146

cemented sediment is visible, This paleosol has

heen identified along Balcanoona and Poontana

Creeks, on the west side of Lake Frome (Figs

1, 15, section 2) where it is developed in

coarser grained sediments.

The Eurinilla Formation is often underlain

by a course croxs-hedded sand or conglomerate,

Yunnamed conglomerate’ usually partly

cemented with hard white lime. The beds are

light pinkish brown, from iron-staining on sand

grains. A typical sequence including this

facies occurs at Lake Pinpa reference section

{section 8, Fig. t5, and Appendix 2). The

basal conglomerate often contains clasts. of

Namba Formation dolomite er Willawortina

Formation carbonate nodules. At Lake Tar-

kurooloo the islands on the lake Aoor (espect-

ally near section 6, Pig. 15, where massive

carbonale cemented pink sands and congio-

merates ate interbedded with the charophyte

limestone), demonstrate the gradation io the

Millyera Formation.

The disconformity between the Millyera and

Eurinilla Formations is exemplified in sections

= and 7, but ig not at all obvious in section 8,

or elsewhere away from the vicinity of Lake

Frome. The relationship can, however, be

observed along the Pasmore River, particu-

larity Where the main Yunta to Flinders Ranges

road crosses. Here, two terraces of secondary

carhonate-cemented conglomerate occur, inter-

bedded with yellowish sands containing green-

ish White carbonate novules at the top. The

nodules. are interpreted as a paleosol, and

occur in similar yellowish sands of the Millyera

Formation in Lake Tarkarooloo. Therefore

the unnamed conglomerate and associated

sands are regarded as Millyera Formation

equivalents.

The red brown silty Eurinilla Formation

with its characteristic soil developed af the

top, infills a valley cut into the ‘unnamed’

conglomerate, and associated sediments. The

whole is cut into Willawortina. Formation

sandy clays. light brown sands of the Coonar-

bine Formation disconformably overlie afl

units at vatinus levels in the landscape,

The relationships between the Eurinilla and

Willawortina Formations is also exhibited in

section 3 at the mouth of the Pasmore River

(Fig, 15 and Appendix 2) where reddish-

brown pebbiy silt with a basal conglomerate

(Burinilla Formation) rests with ‘sharp

erosional disconformuity on pale vreen and red-

browa mottled clay (Willawortina Formation),

R. A. CALLEN & R. H. TEDFORD

On a regional scale the disconformity sur-

face beiween the Eurinilla Formation anc

Coonarhine Formation as flat, but locally, river

valleys are developed,

Within Lake Frome are several islands con-

sisting of up to 10 m thick of coarse well

rounded gypsum sand with minor quartz, and

interbeds of clay pellets. These exhibit the

low angle cross-hedding, lithology and gec-

metry of [unettes described hy Bowler else-

where in Australia (pers. comm, 1974, J. M.

Bowler). The sands rest disconfarmably an

the Millyera Formation indurated clays, and

are tentatively correlated with the Eurinilla

Formation, Similar lunettes flank the eastern

shore of Lake Frome.

COONARBINE FORMATION

Lake Coonarbine,

FROME).

The type section is Jocated at Lake Moko

sevtion 7, Fig. 14 and Appendix 2), mentioned

earlier, The sequence (Fig, 20), resting dis-

conformably on the Eurinilla Formation, con-

sists of three parts—a basal 1-0 m of a red

brown indistinctly Jamibated sand, overlain

by 3,3 m of light brown dune sand (two large

scale cross-bed sets are represented) with a

carbonate soil horizon at the top. This is aver-

lain by 0.6 m of light brown sand with car-

bonate patches and rhizondules, The laminated

sand ai the base of this sequence may be a

distinct unit in its own Tight, since it has

features different from the remainder of the

Coonarbine Formation, Its disconformable re-

Jation with the Burinilla Formation has been

established.

The carbonate soi horizons are much

weaker than those of the Eurinilla Formation

in the same section and at section 5 ¢Fig, 15).

The upper more prominent horizon is corre-

lated with that al the top of section five, The

Coonarbine Formation in this section exhibits

the typical blocky joint pattern, producing 5 x

10 cm columns of sediment (large ped struc-

tures), Land-snail shells occur here, and ul

other widely separated localities, being charac-

teristic of the unit, The uppermost Inyer is

associated with aboriginal artifacts and emu

shel] fragnients. The Formation can be traces

west to the Pasmore River (c.g. section 3)

where it overlies the Eurinilla Formation.

An important supplementary section {sec-

tion 2, Figs 15, 21 and Appendix 2) repre-

senting a coarser facics uf the Coonarbine

Formation of western Lake Frome, is found

(Derivation

Ceonarbine map sheet,

CAINOZOIC ROCK UNITS

in Baleanoona Creek, near the natural gas

pipeline (Air Photo Ref.: S. Aust. Dept. Lands

Svy 394, Artareala Rut 3, Photo No, 0078).

At this site the old land surface on top of the

Lurinilla Formation ts exposed, The overlying

beds of the Coonarbine Formation cansist of

1.70 metres of dark brown, sandy silt, with a

basal pebble bed, moderately poorly sorted,

No bedding plancs are visible, and columnar

ped structure is well-developed.

Immediately downstream the surface of the

Coonarbine Formation is scattered with abori-

ginal artifacts, the colour’ is redder. and Jand

snail shells are present. Upstream, near Mulga

Hore on "Balcanoona", the hasal pebble bed

has 0.3-1 m thivk lenses, cutting into ihe

Eurinilla Formation. Carbonate nodules from

the soils developed in the Eurinilla Formation

are eroded and incorporated into the basal

Coonarbine Formation-

East of Lake Frome the fuviatile facies of

the Coonarbine Formation gives way to

atolian seif dunes, forming the partly indurated

cores of the modern dunes of the southern

Strzelecki Desert, Exposures occur along the

flanks cf the modern dunes. The gypsum

lunettes of the islands have a deposition break

within them, the significance of which is un-

certain: it is likely that the part above the

break correspands to the Coonarbine Forma-

tien,

RELATIONSHIPS BETWEEN

FORMATIONS

The Millyera Formation rests disconform-

ably on the Namba Formation in Lakes Mill-

yera and Tarkarooloo, but its relationship to

the Willawortina Formation is less clear. The

correlation of conglomerates and sands at the

Pasmore River Section with similar facies at

Lake Tarkaroolop has been mentioned, aid

suggests the Millyera Formation conglomerate

equivalent is also disconformable on the Willa-

wortina Formation. The relationship ts similar

to that at the “Wertaloona” section. Further

supporl is derived from the presence of bright

orange to red-brown silt and sand, similar tw

that in the Millycra Formation of section 5,

intertonguing with the conglomerate around

the mouth of Balcanoona Creek,

At the “‘Wertaloona™ section {section 1,

Fig. 3) the dipping scyuence of Willaworting

Formation is overlain with angular uncon+

fommity by a small patch of horizontal con-

glomerate and vellow sand. regarded as

Millyera Formation equivalent. The conglo-

147

merate conmains pebbles of ferruginized

matesinl, derived fromi what was prohably a

widespread siirface, mow exhibited as small

remnants in the same valley. This ferruginiza-

fion & correlate! with that beneath the

Mijlyera Formation at Lake Tarkaroaloo, and

clsewhere. Deformalicn of the Tertiary sc-

quence occurred before deposition of the

Millyera Formation and development of the

ferruginows horizon.

The disconformity between the Eurinilla

Formation and Willawortina Formation can

be seen in clifis along the Pasmore River, The

clearly disconformable relationship between

the Millyera Formation and Eurinilla Forma-

tion is seen in section 5 (Fig. 15), The dis-

conformity 1 less obvious for its equivalent,

the “unnamed conglomerate” of lakes Tar-

karooloo, Pinpa (?units 1 and 2 af section 8,

Fig. 15), aad clsewhere.

Relationship between Eurjnilla and Coonar-

bine Formations can be easily observed (for

example in sections 2, 3, 7 and 9, Fig, 15),

The Coonurbine Formation can be frequently

seen cutting into the Eurinilla Formation, and

the iwo unils usually have contrasting lithology.

The soil carbonate at the top of the Eurinilla

Formation may be completely eroded and

reworked into the younger unit.

Rock relationships are summarized in Fig.

24.

AGE

The Millyera Formation has equivalents at

the southern edge of Lake Callabonna, and

northern end of Lake Frome. ft closely re-

sembles laminated green clays and sands

bearing Diprotoden found in the main part of

fake Cyllubonna. The temporal range of

Diprotedon is Pliocene to fate Pletstocene. A

wood radiocarbon age of >40 000 years BLP.

(Daily 1960) from these beds has lately been

confirmed by unother wood radiocarbon date

of >39 900 years B.P_ (Tedford 1973). At the

mouth of Poontana Creek, on the Lake Frome

—Lake Callabonna confivence, dates from

shells in sands equated with the Millyera

Formation give ages of >33 400 years. BP.

and 35200 +! 200 years B.P_ (GaK-4949,

GukK-4948). This shell materin] has been

allected by younger pedogenesis, converting

them to calcite {assuming the shells were ori-

ginally all aragonite as sare most non-marine

oiolluses). Therefore the dates are minimal,

and the Millyera Formation has an age in

exeess of 34 200 years B.P\, probably >40 000

Las

years BLP, Similar shell beds in a similar strati-

graphic sequence were recorded al Lake Eye,

and gave a date of 39 200 = 1 300 yeurs B.P

(Johns & Ludbrook 1963),

The Eunnilia Formation contains — late.

Pleistocene vertebrate fossils, somewhat dif-

ferent in generic composition to thuse at Lake

Callabonna, The fauna occurs in channels at

the base of the urnl, along Billerow Crock east

of Lake FPinpa

The overlying Coonarbine Formation is

prébably late Pleistocene or eatly Recent.

REGIONAL CORRELATION

Equivalents of Millyera Formation are litle

koown al present, though the sequence de-

scribed imminediately above the Etadunna

Formation in the Madigan Gulf region of

Lake Eyre North is apparently very similar

{King 1956, Ludbrook 1956, Johns & Lud-

brook 1963). The lithological similarity

between the fossiliferous greenish sands con-

taining Coxiella giiesi in Mudigan’s Gulf, and

those in the Millyera Formation of section 5

(Fig, 15) is marked. All these beds are close

to or beyond the limits of radiocarbon daling,

but the closely comparable micro-fauna (in-

cluding Elphidinm spp., Ammonia beceart,

Nonion sp: pers. comm, 4, M. Lindsay 1974),

charophytes and molluscs tend to support

correlation. The Lake Eyre sequence Tests on

the Etsadunna Formatinn, and is overlain by

rocks resembling the Ticrari Formation,

The Eurinilla Formation closely resembles

the Tirati Formation of the Lake Eyre Basin,

in lithology, stratigraphic position and topo-

graphic expression, Vertebrate faunas in basal

Eurinilla Kormation channels indicate equiva-

lence with the youngest Kutapiri Sand (Stirton

et al. 1961) of the same basin.

Other possible equivalents are indicated in

Table 4. Vhe Poorska Formation (Firman

1956a) supposedly resis un Telford Gravel

(Firman 1963) on the west side of the Flin-

ders Ranges, and is overlain by the Lake

Torrens Formation (Williams & Polach 1971).

The unnamed conglomerate equivalent of the

Millyera Fermation lithologically resembles

the Telford Gravel at Telford open cut, Leigh

Creck. The Eurinilla Formation, lithologically

resembling the Lake Torrens Formation, aver-

lies the Millyera Formation, and ts in ttm

overlain by the Coonarbine Formation. The

lane is similar to the Thomson Creek Forma-

tion GF Willianys & Polach. There also are

R. A. CALLEN & R. H. TEDFORD

similarities in the calcareous soil horizans of

each, in the same geomorphic situation.

The Pooraka Formation, Telford Gravel and

“unnamed conglomerate” of Lake Prome area

are probably equivalents. Jf has been suggested

by Firman (1971)* that the Telford Gravel is

eguivalent to the whole of the Tirari Forma-

tion (Eurinilla Formation correlative), but

this cannot be the ease in the Lake Frorne

area, The youngest profable equivalents here

are the convlomerate at the base of the

Burinilla Formation, and the most likely corre-

lative the “unnamed conglumerate” equivalent

of thé Millyera Formation.

The unit mapped as Pooraka Formation on

COPLEY (Coats 1973) ts Coonarbine Forma-

tion. During mapping COPLEY, Callen &

Williams (in Coats 1973) recognized a unit

of reddish brown sand and cobbles which

covered most of the surface of the high level

plains flanking the eastern Flinders) Ranges,

The unit was Jater named the Afrowie Forma.

tion by Coats {1973}: subsequently mapping

for FROME has shown it is probably partly

equivalent to the Coonarbine Formation. The

two units beth contain Jand snuil shells, and

appear to grade laterally into one another at

the bresk in slope at the basc of fow hills

south of “Wertaloona,” However. Coats secms

to include some vounger and older gravels in

his definition, with %disconformable relation-

ships.

ENVIRONMENT

The Millyera Formation constitutes three

facies groups: the most typical and widespread

ate Yhe laminated ostracode clay and chara

phyte limestones (Fig. 18), with associated

charophyte eogonia-bearing fine sand. Fine

lamination, ostracodes, and distribution of

sediment, indicate they are undoubtedly of

lacustrine origin, The fine sands are well

rounded and smooth and mav therefore be

aeolian, haying been blown inte the lake, or

carried by floods. Drying of the lake is indi-

cated by the charophyte limestone and equiva-

lent gypsum lamellae (Figs 16, 18; cf. Reeves

1968, p. 57, 58). Similar madern calcareous

algal deposits (Fig. 19), grading to rippled

gypsum crusts, arc present in Lake Kuturu’.

Waves acting on the very shallow water badies

break vp the filaments and orient them in

crescent like ripples, sometimes resembling the

oriented structures in thelr fossil equivalents.

The gypsum Jaminac may have hotryaidul

surfases that are reminiscent of similar forms

149

SeUIW 4O JUaWjiodag “ys

vaya va 10¥—PvZ

—e— —t— shol> puo |

SaNVS |

NOOMNdWYW

AQYNSTLYND

AVa¥d

Ol AaVILd1

i1¥1

33} Dia WO] Buo> , Z : |

SU St Cl NOILYWYOS VNILIOMVTIM

pawouuy |

O-L TWAVID |

({luA Bulloag

uapojoidig)

"Ainby NOILYWHOS

“WaSATIIW

ONYS ldld¥ivy Pus

NO|LYWaOd ldVaiL

(un Buioag

—Psiq |!s8o4)

juajoainby

NOllyvwaod

INIad¥NOOD

CAINOZOIC ROCK UNITS

BuVogD||D> 2x07 8143 407 7961

("Wor 'siad ‘E76 1)

a4¥Osd3L

“y2 48 NOLMILS

TOsOAlWd YNVLVIIUIM

NOILVWYO4

SNaddOL IVT

1OsSOIIWd

YNad Low

(j40d JaMoj)

NOILYWao4

4434S NOSWOHL

1OSOIWd

VNOOOYN

{od uaddn) -4

43D NOSWOHL

saBuoy

SUPPUlfy Wiad A

‘1261 ‘HDV1Od

PYO SWYITTIM

dyOsTdL

T9A¥aID GYOITIL

MOS Vard¥a

NOILVwaO4d

VAVYOOd

WOS Yavnvee

WOs AV3A07

Os Vavevd

ONY ALadDdAD

syisodap Builds punhow

NOILY¥wacd

Va¥aoOd

NOllYWwao4

NOILVWaO4

WAVYOOd

WOS AVTIAOI

_ -aIMOxay

6— =~ AVYD

YNNOBVTI¥>

GNV¥S NOSdWIs

uIsog asAQ 2407

‘saBuoy

SJEPUl|) UsBISaAA “Nh

"NY WHld

jeays dow

000 OS2*1 AIIdOD

“JB 49 SivVOD

NOlLWWwaod

VaFATIIW

aLaysW> ; :

NOILYWdO4

VINIGNG

(MIDjsaaun aaug

alvwaWOISNOD

OWYNNN

Gig ayydosGyoug)

YOrOW oO, By) uing |Ssoq jo

JLFYDIVD PY FL3¥DdAD

UOZIIOY [105 snoasosj0> jisso4

sajjaun| wosdAB sapnjour

INDI

NOLVWHOI INIEYYNOOD

-IN3D015131d

UOZIVOY |[Os sMOBIOD}O> jisso4

POojpag woays

syisedsp 9904

saunp usapow

aadod siyy

QdOsdd1 ® NAlIVWS

SNISVS Y3HLO ONV SL3SHS d¥W LNADVray - LNZIVAINGA GaLsa99Nns

SLINM YI9NNOA - LYVH) NOILVTIAYO)

yay

3WOYs AVI

¥ A18V1

180

on the sufface of modern Lake Frome, pro-

duced by crystallration pressure bockling the

surficial crusts,

The second facies group is the channel facies

(Fig, 17), of conglomerate/sand which ex-

hibits Features of meandering streams of large

size containing bed-forms of slightly crescent-

shaped aqueous dunes. The streams carried

pebbles from the Olary Ranges, and eroded

valleys into the Namba Formation, These

deposits are lateral equivalents of the “un-

named conglomerate” which is so extensive

along the Siccus—Pasmore River System.

The third facies group are the greenish fos-

siliferous sanis, which (Section S, Fig, 15}

are cross-bedded on a small to medium scale,

and contain shell beds and fish vertebrate.

Similar shelly are also abundance in a narrow

zone slong Billerog Creck between Lakes

Kuturu and Tarkarooloo. These deposits ate

interpreted us shoreline facies of the Pleisto-

cene Lake Frome.

These sediments, and equivalents at the

northern end of Lake Frome, contain the

foraminiferal assemblage mentioned earlier

(p. 147). Similar species were also recorded by

Ludbrook (Ker 1966, p. 94) in equivalent

strata in McKenzie Bore, 7,5 km sauih south-

eust of section 5, The presence of several

species of foraminifera over a wide area in the

same scdiments can be explained in terms

of Ludrook’s (1965) hypothesis of transport

to salt lakes on the feet of seubirds, with

subsequent survival for a period, The species

R. A, CALLEN & R H, TEDFORD

present ate mostly Rotaliina with a wide

salinity tolerance, and diversity is low. Such a

situation ts typical of inland saline lakes (Resig

1974), where foraminifera have been intro-

duced by some dispersal mechanism from

coastal areas. Species such as Ammonia bec-

carti are common in these environments.

Although the assemblages found at Lake Eyre

and Lake Frome are considerably different in

content from those listed in Table 4 of Resig’s

paper (e.g. Nonion spp. are not recorded,

though common at Lakes Frome and Eyre)

this does not detract from the dispersal hypo-

thesis because each locality cited in her paper

has high endemism. The Coorong area con-

tains « similar assemblage (pers. comm. J. M.

Lindsay 1975), though its low diversity is pro-

bably the result of high salinity, even though i

has a connection with the sea.

Another explanation is that there was a

distant connection to the sea, implying a high

sta level during the Pleistocene prior ta 40 600

years BLP.

The detrital component of the facustrine

Millyera Formation sediments were brought to

the ancestral Lake Frome by large braided

streams with a pebble bed load (“unnamed

conglomerate”) approximately following the

channels of present day watercourses such as

the Pasmore-Sictus River system, and the

Luke Tarkarooloo-Billeroo Creek system. They

were much more extensive than their modern

counterparts. The clasts. indicate 4 provenante

similar to the modern streams, li the Olary

Figs 16-19. Younger Units. Structures in Millyera Formation.

Fiz, 16, Millyera Formation. Laminated tipple-murked gypsum and clay (Fig. 15, section 5). Scale 30

cm.

Pig.

17, Plan view of cross-stratificd channel sand in Millyera Formation channel facies, bed of Lake

Tarkacooloo near Section 6, Fig, JS, Approximates Pi cross-stratification. Current direction

(arrowed) is te north. Hammer handle 25 cm long. Lominace emphasized Sy inking.

18, Algal tubules showine rough onentation. Same jocality as Fig. 22. Scale in cm.

. 19. Modern, calcarcous charophyte algal flaments, Lake Kuturu, showing crode orientation. Thin

crust of gypsum {G) in upper central part of photograph.

Figs 20-23 Oitecrop,

20. Upper part of section 7, Fig. 15, showing dune sand facics of Coonarbine Formation (1wo

upper benches), basal laminated sand (bench Just above contact), and tipper part of Furinilla

utmation with calcareous paleosol (just below contact).

. - Section 2, Fig. 15, Columnar-structured sand of Coonarbine Formation overlying Enrinilla

Formation. Surface in foreground shows carhonute patches of paleosol, and represents the pre-

Coonarbine Formation land surface slightly modified by present erosion. Scale 30 cm.

2. 22. Coonasbine Formation sand with columnar jointing, overlying Millyera Formation which in tum

avetlies Nainba Formation. Millyera Formation shows wpper algal limestones und lower mias-

sive sandy Jimestone (prominent benches) with intervening clayey sand. Lake Tarkaronloa,

near Coombes Bore- Scale 30 cm,

Rig. 23. Section 8. Fig. 15 Cower part}. Coonarbine Formation, disconformably overlying Eurinilla

Formution which has its opper surface cemented with secondary gypsum (prominent hench)-

Black vlay of Namba Formation at base (30 cm) scale crosses contuct)-

COONARBINE

CAINOZOIC ROCK UNITS

a Ls gio -é

a “e J * ‘i

EURINILDA’ Be

152

Ranges and southern Flinders Ranges. Large

straight-crested aqueous dunes typified the

streams with coarse sandy and pebbly bed

loads, whereas crescentic dunes characterized

the streams with a finer sand-bed load, The

eastern shore of Lake Frome was estimated to

be about [0 km further cast than al present,

The Eurinilla Formation contains channel

deposits, exemplified by coarse sand with

parting lineation and cross-bedding, in troughs

and point bar deposits along Billeroo Creek,

The meandering form of these channels can

sometimes be seen on aerial photographs. The

pebbles have sources in the Flinders. Ranges

or Olary Ranges, or have been eroded from

the underlying Tertiary units. Flood-plam de-

posits are represented by the finer facies,

which is sometimes laminated, The initially

fluviatile phase (basal coarse grained sands)

gave way fo a more complex environment with

finer fluviatile deposiis and large scale cross-

bedded aeolian deposits, including huge gypsum

lunettes along the south western shore and on

the islands. Some possible Joess (massive silt

and very fine sands) is present, These sedi-

ments transgressed over the older lake deposits

of the Millyera Formation, The ancient Lake

Frome therefore decreased in size in medial

Pleistocene times, being somewhat smaller than

at presenl,

The plains of this essentially fiuviatile en-

vironment were inhabited by large marsupials

(Dipratedon sp., Procaptadan geliah, Sthenu-

rus sp. and Macropus sp.) Rivers followed

EURINILLA FORMATION

UNNAMED CONGLOMERATE

WILLAWORTINA FORMATION

Eocene

2 Paleocene

S1O970

COONARBINE FORMATION

EYRE FORMATION

R.A.CALLEN

R. A, CALLEN & K, H, TEDFORD

approximately the same coarses as the present

day drainage. The distribution of lunettes indi-

cates a dominant wind direction from the west

and a strong westerly component still charac-

terizes this region.

The overlying Coonarbine Formation in

cludes fuviatile braided stream environments

west of Lake Frome, and dominantly aeolian

¢ast of the lake. The fuviatile sediments have

less defined channels than the Eurinilla Forma-

tion, pebble sheets being more common. East

of Lake Frome Jongitudinal dunes were deve-

loped, and another minor phase of gypsum

lunettes built up along the lake shore. Land

snails probably lived around water holes.

Conclusions

The new rock units in the Lake Frome area

record a history of intermittent deposition

through Miocene to late Pleistocene-Recent

times, During this interval the extensive rivers,

lakes and possibly estuarine environments of

the Miocene Namba Formation drained areas

of low relief in a climate of high rainfall, and

of higher annual temperature than the same

latitude today, At times, seasonal dry periods

became a part of the weather pattern. A con-

nection with the sea was established at some

stage, probably to the Murray Basin, Some

conflicting climatic evidence is partly resolved

by applying the continental Jessivage hypo-

thesis (Millot 1964, as modified by Wiersma

1970) in relation to the smectite—dolomite—

palygorskite mineral suite, Thus warm lem-

Member 2

Member |

NAMBA FORMATION

S.A. Department of Mines

Fig, 24, Cuinozoic rock stratigraphic relationships, Lake Frame.

CAINOZOIC ROCK UNITS

perale to subtropical conditions prevailed, with

savannah landscape, and gallery forests around

the Jatge permanent streams and lakes,

Uplift of the Flinders Ranges securred at

the earliest during late Miocene times, con-

tinved through the Pliocene intermittently into

the Quaternary, and is still proceeding at

present, Prior to this, at least during the

Cainozoic, the Flinders Ranges were virtually

non-existent. The sediments deposited during

the Pliocene-early Pleistocene Epochs record

the change from the earlier Miocene palaeo-

gcography to the very different landscape

approximating that of the present. Lakes and

swamps during Tertiary times disappeared

dunop the Pleistocene, as tectonism and

Climalic change altered the depositional

regime. Drainage resembled that of the present

during the late Pleistocene, indicating the basin

was approaching its present configuration,

The Millyera Fotmation indicates active de-

position on a playa lake somewhat larger than

ihe present. The changing character of the

sediments from Millyera to Coonarbine For-

mation suggests overall increasing aridity, pro-

bably seasonally distributed during Eurinilla

Formation limes, as exemplified by the forma-

tion of the gypsum Iwnettes. Marked climatic

Muctuations were superposed on this overall

climatic trend. Uplift of the ranges continued,

153

alternating with periods of stability during

which soils developed.

Rock relationships are summarized in Fig.

24.

Acknowledgments

This paper is published with the permission

of the Director of Mines, South Australia

Drafting and typing facililics were provided

by the South Australian Department of Mines,

Participation of R.H,T. was supported by

National Science Foundation Grant GB-

b8273X.

The authors also wish to acknowledge dis-

cussion and helpful criticism by Mesrss J. B.

Firman, W. K. Harris and J. M. Lindsay of

ihe South Australian Department of Mines,

and by Dr B. Daily of the University of

Adelaide. The project was initiated by Me

B. P. Thomson of the South Australian Depart-

ment of Mines.

Radiocarbon dates were supplied by K.

Kigoshi, Gakushuin University, Tokyo,

The following companies provided core

material and cuttings: Mines Administration

Pty Ltd, and Mr J, Andrus of Western Nuclear

Aust. Pty Ltd is especially thanked for arrang-

ing WC2 bore to be donated for use asa type

section.

The skilful drafting of Mr Bruce Thomas

of the South Australizn Department of Mines

i acknowledged,

References

ALLEN, J. R. Lb. (1963)—The classification of

cross-stratified units, with notes on thelr ori

gin. Sedimentol. 2, 93-114.

Benton, Y. K., Bopenneimer, W,, & Herrer, L.

(1963)—Clay minerals mm Negev sediments.

J. sedim, Petrol. 33, 874-903,

Bryxs, P. J, (1972) —Late Cainozoic uplift of the

Ediacara Range, South Australia, Proc. As-

erelec. Inst. Min. Metall. 243 (Sep. 1972) 47.

BUISSENBACH, B, (1954),—Geology of alluvial

fans in semirarid regions. Bull. zeal. Sov. Am,

6S, 175-190,

Brewes, R. 41964),.—"Fabric and Mimeral Analy-

sig of Soils." (Wiley: New York, London,

Sydney) 470 pp,

Baown, A. Y. 1, (1884) —Report of the Govern-

ment Geologist. Parl. Pap, 3. Aust. 35, 1-20.

Catten, R. A. (1975)—FROME geological

sheet S1154-10. Geol. Atlas Ser, geol. Surv.

S. Aust.

CaunEnN, R, A. (1976).—Lake Frome area—

regional geology, Tertiary Stratigraphy and

uranium localization. Jn Knight. C. L. (Ed.),

"Economic Geology of Australia and Papua

New Guinea”, Vol. I. Metals, Monagr. Awy-

tralas, Inst, Min, Met, 803-808.

Coats, R, P. (1973)—COPLEY map sheet, Exe

Planatory Notes 6-250 000 Geol, Atlas Series,

pect SH54-9, Geol. Surv. S. Aust. (Ade-

aide),

Daice, B, (1956).—The Cambrian in South Aus

tralia, Proc. 20h int. geal, Cong, (Mexica)

2, 91-142.

Datty, B. (1960)—Report of the Curator of Fos-

sils and Minerals, Repr 8. Aust, Mus, Board

1958/59, p, 12 (Adelaide),

Denny, C. S. (1967).—Fans and pediments. Aint

J. Sei, 265, 81-105.

Devereux, I. (1968)—Oxygen isotope paleotem-

peratures from the Tertiary of New Zealand.

Tuatara 16(1 4, 4144,

Finman, §, B. (7963),—Quaternary geological

events near Swan Reach in the Murray Basin,

South Australia. Qaert, Geal, Netes, geo,

Surv. S. Aust, 5.

Firman, J. B. (1964).—The Bakara Soil and

other stratigraphic units of the late Cainozode

Ave in the Murray Basin, South Australia,

Quart, geol. Netes, geal. Sued, §, Aust. 10,

-5,

Firman, J. B, (1966a).—Stratigraphic units of

Late Cainozoic Age in the Adelaide Plains

Basin, South Australia, Quart. geol. Notes,

geal. Surv. S, Aust. 17, 6-8.

154 R, A. CALLEN & R H. TEDFORD

Finman, J, B, (1966b).--Stratigraphy of the

Chowilla area in the Murray Basin, @uart.

geol. Notes, peol. Surv. S. Aust. 20, 3-7.

Firman, J. B, (19672)—Late Cainozoic strati-

graphic units, Quart. geol, Notes, geol, Surv,

S, Aust. 22, 4-8,

Fiemam, J. B. (1967b).—Siratigraphy of Late

Cainozoie deposits in South Australia. Trans.

R, Soc S. Aust, 9, 165-178,

Firman, §, B, (1969),—The Quaternary Period:

In Parkin, L. W. “Handbook of South Aus-

tralian Geology”. (Govt Printer: Adelaide).

Firman, J, B, (1970),—Late Cainoznic strati-

graphic unils in the Great Artesian Basin,

South Australia, Qvart, geal, Notes, geet,

Sarv. S. Aust. 36, 1-4.

Foik, R. L.. ANprews, P, 8, & Lewis, D. W-

(1970) — Detrital sedimentary rock classifi-

cation and nomenclature for use in New Zea-

Jand, N.Z. J. Geal, Geaphys. 13, 937-948,

Frevret, P. (1971}.——Paleosols residuels et paleo-

sols alluviaux hydromorphes dans le Cretace

superisur el ['Eocene basal en Languedoc.

Revue Geapgr, phys, Geol. dyn. 2, 245-288.

FREIDMAN, G- . AmreL, A. J, Braun, M. &

Mitsera, D. 8S. 41973),—Generntion of car-

bonate particles and laminites io algal mats—

example from sea-marginal hypersaline pool,

Gulf of Aqaba, Red Sea, Bull, Am. Ass. Per-

rol. Geol. 57, 541-557.

Grr, E. D. (1968) —Onygen isotope palaeatem-

perature determinations from Victoria, Aus-

tralia, Teotare 16¢L), 56-61.

Hayes, BE, & Frakes, 1. A. (1973).—Leg 28,

deep-sea drilling in ihe southern ocean, Geo-

limes 18, 19-24.

Howcnrn, W. (1909),—Description of an old lake

urea in Pekina Creek and its relation to

Recent geological changes. Trans. R- Soc. S-

Aust, 33, 253-261,

HowcHin, W. (1913}—The ¢volution of the

physiographicol features of South Australia.

Tn: Australasian Assoc. Adv. Sct. 14, p. 168:

Hall, T, S. (Ed,

Jack, R. L. (1930).—CGreological structure and

ather factors in relation to underground water

supply in portions of South Australia: Bull.

geal. Surv, 3. Aust. Wa.

Jenxins, D., G. [1968).—Planktonic foramini-

ferida as indicators of New Zealand Tertiary

paleotemperatures. Tuatara 16( 1), 32-37.

Jessup, R. W. & Norris, Ri M. (1971).—Caino-

zoic stratigraphy of the Lake Eyre Basin and

part of the arid region lying 10 the south, J.

geol. Soc. Aust. 18, 303-333.

Jouss, R. K, & Luonroox, N, H, (1963),—Inves-

ligation of Lake Eyre Parts 1 and If Repr.

(nvest,, geoi. Surv. §. durt. 24.

Kenny, EB. J. (1934).—Darling districi—a_ peo-

logical reconnaissance with speciul reference

io the resources of subsurface water, Mineral

Resour. N.S.W. 36.

Ker, D. S (1966)—The hydrology of the Frome

Embayment in South Australia: Rept. Invest.,

peal. Surv,, §. Aust. 27,

King. D. (1956).—The Quatetnary stratigraphic

record at Lake Fyre North and the evolution

of existing topographic forms. Trans. R, Soc,

¥. Awst. 79, 93-103.

Krinstey, D. B, Woo, C. C. & Sroerrz, G. E.

(196%) —Geologic characteristics seven Aws-

Irslian Playas: Ju Neal, J. T. (Bd.), “Playa

surface morphology; miscellaneous investiga-

tions’ OS. Air Force Cambridge res. Lab.

Environ. Res. Pap, 283, $9-103.

Linnsay, J, M. (1969)—Cainozoic Foraminifera

and stratigraphy of the Adelaide Plains Sub.

basin, South Australia, Bull, geal Surv. §.

Aust. 42.

Linpsay, J, M, & Srervearn, R. G. (1966)—

Muono Para Clay Member. Qwart. geal.

Notes, geol. Surv, S. Aust. 19, 7-11.

Luperoox, N. H, (1956}—RMicrofossils from

Pleistocene to Recent deposits, Lake Eyre,

South Australia. Trans. R. Soc. S. Auss. 79,

37-45.

TLuprroox, N. H, (1965) —Occurfence of fora-

minifera in salt lakes. Quart, geol, Notes.

geo! Surv. §. Aus 14, 6-7.

McLean, 8. A., ALLEN, B. L, & Craic, J. R-

{1972),—The occurrence of sepiolite and

altapulgile on the southern high plains. Clays

Clay Miner, 20, 143-149,

MEESTER, T. DE (197L)—Highly calcareous lacu-

strine soils in the Great Konya Basin, Turkey.

Versl. landbouwk. Onderz (Agr. Rept) 752.

Mrvzaor, G. (1964)—Trans. Fanranr, W. R, &

Paquet, H. (1970),—“Geology of Clays".

(Springer-Verlag: New York, Heidelberg,

Berlin, Paris; Chapman & Hall: London.)

Muis.er, G., Tron, G, & Forstner, U, €1972).—

Formation and diagenesis of inorganic Ca-Mg

carbonates in the lacustrine environment.

Naterwissenschalfer $9, 158-164,

Muaray, J. W, (1968).—Living foraminifera of

fngoons and estuaries. Micropaleentalegy

14, 435-455.

O'Driscoit, E. P. D, (1956),—The hydrology of

the Willochra Basin. Repl Invest., peel. Surv,

S, Aust, 7, 58 pp.

Reeves, C. C. (1968).—“Introduction to Paleo-

limnologsy", Developments in Sedimentology

Hl (tElsevier: Amsterdam, London, New

York).

Resic, J. M. (1974)—Recent foraminifera fron

& landlocked Hawaiian Jake. J. Foraminer,

Res. 4, 47-60.

Sriwrn, A. R. C. (1860),—Geological notes an

a journey in Somh Australia from Cape Jer-

vis fo Mt Sorlc. Parl. Pap. 5. Aust. 20.

Sincrr, A,, Gar, Mo & Bantn, A, (1972) —Clay

ming¢rals in recent sediticots of Lake Kin-

neret (Tiberias), Israel, Sediments. Geol, 8,

289-308.

Smuen, A. & Norurso, H. K. (1974). —Pedogenic

palygorskite aceurrences in Austtalia, An,

Miner. 59, 508-517.

Staton, KR. A,, Tevrorp, R, H, & Mincer, A, H,

(1961)—Cenozoic stratigraphy and verte-

hrate palaeontology of the Tirari Desert,

ha Australia, Rec, 8. Aust. Mus. 14, 19-

Smeaton, R. A.. Tepronn. R. H. & Wooprurne.

and fauna from the Tirati Desert, South Aus-

tralia, Ree. S. Airst, Mus. W, 85-93.

CAINOZOIC ROCK UNITS 155

SUTHFRLAND, F, L, Green, D. GC. & Wart,

B. W. (1973),—Age of the Great Lake

Rasales, Tasosania, in relation to Australian

Cainazoie volcanism, J, geal, Soc, Ansi. 20,

5-93.

Tepeoap, R. H. (1973)—The Dipretodons of

Lake Callabonna. Aust. Nat. Hist. 17, 349-

354.

Top, D. H. (1968).—Palaeoclimatology and the

relative stability of feldspar minerals under

almospheric condition. J, sedim, Petrol, 38,

§32-844.

VON DER BorcH, C. C. (1965)—The distribution

and preliminary geochemistry ef modern car-

bonste sediments of the Coorong area, South

Australia. Geochim, Cosmochim, Acta 29,

781-799,

VON DER Boacn, C. C., Lock, D. E., & Sciwener,

D. (1975),—Grountwater formation of dola-

mile in the Coorong region of South Aus-

tralia. Geology 3, 283-285.

WeLiMan, P., McEcamny, N. W.. & MeDoucart,

[. (1969)—On the Polarwander path for

Australia during the Cenozoic. Geophys, JR,

aatr. Sov. 19, 371-3958.

Wirrsma, J. (1970).—Provenance, genesis and

palcogeographical implications of micro-

Minerals occursing in sedimentary rocks of

the Jordan Valley area. Pr-Fys. Geogr. Bo.

Univ. Amsterdam 15,

WituiaMs, G. EB, & Potacnh, H. A. (1971)—

Radiocarbon dating of arid zone calcareous

paleosols, Ryil. geal, Soc. Am. 82, 3069-3086,

Woprner, H. -(1974),—Post-Rocene History anc

stratigraphy of northeastern South Australia,

Trans. R. Sac. S. Aust, 98, 1-12.

Worerner, H., CALLEN, R. A,, & Hamas, W. K.

(1974),—Lower Tertiary Eyre Formation of

the southwestern Great Artesian Basin, J.

geol. Sov. Aust. 21, 17-52.

Appendix I

OLDER CAINOZOIC UNITS

Fig. 3, sections 1, 10, IL, 12. Fig. 14, section 13

NAMBA FORMATION TYPE SECTION

SADM Yalkalpo No. 1 Sttatigraphic Bore. Fig. 3 Section 12

COONARBINE FORMATION

2.00. SAND. medium grained, strong reddish brown (2.5YR4/6). Subrounded grains. Rec-

tangular joinc pattern und carbonate cylindroids in upper part.

— Disconformity —

NAMBA FORMATION

Unie 18 140 CLAY; slightly silty, with scattered gypsum spois, Light grey (N.4+) with moderate yel-

3.40 lowish orange (([0YR5/6) patches, Sharp upper contact.

Unit 17. 1.35. Interbedded SAND and CLAY overlain by SILT. Sand is very fine grained, laminated

4.75 and small scale cross laminated, a3 are the silt beds at the top of the unit, Lower beds. have

structure destroyed by secondary gypsilicution. Contacts between sand and clay beds are sharp

and flat. Sand-filled shrinkage cracks extend down from the wavy irregular upper contact.

Unit 16. 645. SILTY CLAY. Poorly sorted, diffusely laminated. Two silt beds with gradational to

12,00 sharp contacts near the top, lower 7 m burrowed and bioturbated with wavy irregular upper

bedding plane separating it from the remuinder of the sequence. Irregular shiny fracture planes

(crumbly texture) and gypsum nodules developed in upper (part. Light to mederale olive grey

(S¥4/2-SYS/1, 5-SY6/1). Mottled black and yellow (S5Y4/0.5) in lower silt.

0.60. No recovery.

0.20, As before. Shrinkage cracks filled with sand extend down from upper unit—wavy

irregular contact, Light olive grey (SV5/2).

Unit 1%. 0,20, SILT, Laminated, very fine cross-laminated, burrowed in par Very pale yellowish grey

12.20 (NI to 5¥9/1), Upper contact sharp and flat. burrowed.

Unit 14. 3,00 SILTY CLAY. Intraformationally brecciated. Borrowed and bioturbated in lower half,

£5.20 upper half with angular clay clasts and slump structures, Upper contact sharp, flat.

Unit 13, 1.70 CLAY SILT and CLAY, Poorly sorted, Clay beds near centre. Rurrowed at top, intra-

17.30 formational breccialion common. Pale to light olive clay (10Y6/2 to 6/4), sill pale grey to

pale yellowish grey (SY8/1 to N7), Upper contact sharp and flat.

0.40, SAND and SILT, interbedded, weakly laminated, wavy ‘sharp contacts between inter-

beds. Sand is very fine, very angular (quartz crystal faces—overgrawths on rounded grains,

contacts between overgrowth and erain visible),

Unit 12. 1.05. No recovery.

19.89 1.24. SAND, grading up to CLAY SILT and SILT. Sand hus small-scale cross lamination with

heavy mineral luminae.

Unit 11. 2.05, CLAY, black as below, Contact with overlying bed sharp and fiat.

21,52 1.13. No recovery,

} 0.75. SAND, grading up to CLAYEY SILT. Sand very fine, wavy lamination, Colour SY4/4,

Unit 10. 0.20, STILT and CLAY, Intertaminated, flame structures on contacts. Light olive grey (Clay

26,82 SY6/4, Silt NO). Contact with ynit 11 sharp and flar.

2,30, SILT. lower bed laminated .and cross-bedded, with scattered burrows.

result of bioturbation and extensive burrawing. Upper contact wavy and irreg tar,

9.30. CLAY, laminated, colour NO, 4¥4/2, With CHRLONIA scute at 24,21 mm. Upper con-

tact sharp, with flame structures.

156 R- A, CALLEN & R. H, TEDFORD

1.40, SAND, fine grained in lower 1/5 grading up to !aminated and burrowed light grey

(NS) SLLT. Sand grains smooth and shiny, angular, some well rounded and frosted, many

with crystal faces and re-entrants. Upper contact wavy und irregular. Grading down to - .

1.10, SIL], laminated and small scale cross-hedded, with scattered burrows:

Units. 3.20, CLAY. black (N1) mottled light olive brown (5Y5/6), with sand patchcs and other fea

35.52 tures as before, Contact with sand lower in unit gradational. Large sand grains in the patches

are polished, rounded to well rounded, smaller graims being angular to subangular with over-

growths{?). Some of the larger grains show rounded crystul forms.

0.35, Ne recovery,

3,60. SAND, us below. Calcite patches and. very coarse mica common. Polymodal, poorly

sorted overall, Small sizes angular, coarse are rounded, some doubly terminated crystals. In-

terbedded light grey (N7) clay in centre of unit. Becoming well sorted and fine grained at. top

with mixed well rounded and angular grains, Obscure small scale cross bedding and Jamination,

0.60, No recovery,

095, SAND, Coarse to medium grained, slightly calcareous. Large grains polished, others with

crystal faces (overgrowths?) which give siepped shiny surtaces. Many grains shew original

elongate quartz prism shape. :

Uai.8 0.70. Alternating SAND and CLAY, Sand very fine grained with small scale trough cross-

36.22 lamination. Clay olive to medium grey (SY5/0.5).

Unt? oan. Clay as below, Sharp wavy upper cantact,

39.22 1.25. No recovery.

145, Alternating SAND-CLAY fining upwards sequences, Several thin beds, beginning with

very fine grained sand al base. Sharp !lat contacts, grading to black (NZ, SYR2/1) clay with

orange brown specks and sand patches in top 21cm,

Unit6 0.94, SANDY SILT. grading up to CLAY. Hlack (N2) clay as for unit 5, with fine sand

40.79 patches, 1/3 of sequence. Sharp wavy upper contact.

0.58, No recovery,

0.05..SILT with slump structures.

Unit 5. 6.55 SAND, SIT-T and CLAY alternating in lower 1/3, grading to CI.AY at top. Lithology a3

47.44 for unit 4. Obscurely laminated in lower part, lenticular bedding, Light olive grey (SY6/1)

with oxidised brawn patches (LOY R67). Calcareous at transition (25 cm) to dark clay. Sand

distributed in vertical strevks and patches through the dark olive clay (5Y5/1). Irregular

wavy upper contact, - A

Unitd. 3.30. SILT, grading up to CLAY. Lower 1/3 laminated silty and clay with very-small to small

50.84 scale crass bedding st base, some burrows. Pale yellowish grey (S¥9/1) and pale olive

(JOY6/24. Grades rapidly into sandy ¢lay with vertically orténted structure and lime streaks,

yellowish grey to pale olive (3V7/2 to 9Y8/2), This grades fo tough black (NI) clay with

erange brown dendrite mottling and patches of fine sand. A thin brown band of ton oxides

is present. Upper contael wavy, irregular.

Unit, 3. 1.20. SAND, grading to CALCARHOUS CLAY. Lower 40 em very fine grained, loosely

52.04 cemented by guartz overgrowlhs (original grains rounded) calesreaus at base, Heavy minerals

1%, Dusky yellow (SY4/1), Clay is olive crey (5¥4/1) and has white vertically oriented

streaks und sheets of carbonate, Contact with overlying unir is sharp and wavy.

Unit 2. 1.94. No recovery.

56.60 0,22. SAND, very fine grained as for bage unit, Contact with underlying clay irregulai, with

mixing.

1.10, CLAY, as for base unit 1, silty top, with moderate yellow green clay patches which be-

come dull on exposure (7GY4/1, 5Y5/2, 5¥4/2). Obscure lamination.

1,00. No recovery. P . .

0.20. SAND, very fine gfained, scattered medium, polished grains, Opaques common. Sone

palches modenue yellow green clay (7GY4/1-5Y4/2).

Unit |, 1.20 CLAY, waxy lustre on curved ipregular fractures. Rare angular white carbonate lumps

and streaks. Olive grey (SY¥S/2Z)-

WNAMBA FORMATION

Supplementary, Section Outcrop, West Sule Lake ‘Tarkarooloo, Section 13, Fig. 14

Section 13A. north of the northern track crossing Luke Tarkarvolao.

COONARBINE FORMATION

bid, SAND. Very culeareous, with numerous 1-5 cm dolomite nodules and ferruginous sandstote

lumps (reworked from Namba Formation). SYR8/5-

— Disconfermity —

EVRINILLA FORMATION

2.40. CLAY-SILT-SAND. Silt (dominant) to fine grained sind, poorly sorted, Lower If om moderately

sorte medium grained sand. Up to §0% reworked dolomite madules (pisolitic) of granule to pebble

sixe in lower part, 72 x 5 cm maxianim size, and little evidence for abrasion, Upper 10-70 em with gyp-

sim nodules and curbonute patches. Green-bluck terrruginous and manganiferous stain, Olhcrwise lel

red-brown 2.5YRS/6,

| — Desvonturmity —

NAMBA FORMATION

3.85, CLAY, GYPSUM, Slightly silty clay, very hard, ileht weight. crumbly with greasy lustre, Colour

CAINOZOIC ROCK UNITS 137

SYS/2 ta 9/1, mottled red-brown, yellow brown and black, specked with while gypsum flour, 80 cm

hetizon of caulilowcrokt gypsum nodules 1,10 m from top. Nodules 0.25~0.50 om dian. with clay

Cores, nt associated with any porosity change in host sediment.

1.25. DOLOMITE, CLAY. Lower dolomite 6.60 m clay 0.50 m upper dolomite 0.15 m,. Lower dolo-

mile nodulue (intraformatianal conglamerale} al lop, manganese dendrites (hraughout. White, very fine

grained, with 5% angular silt, Upper dolomite contains natracades. Clay a3 above,

1.95. CLAY. S% angular silt to very fine sand, grading to sand at base and dolomtite at top. Very hard,

jight, dry and crumbly, with manganese stain.

145. SILT. Sift and very fine sand, becurning clayey at lop. Bioturbuted ul lop, 2-3 cm burrows, other:

wise finely Laminated, Minor bright green clay faminae, Top 40 cm impregnated with gypsum, mortied

orange browse Very light grey to white overall, Lower contact sharp, far.

70. CLAYSILT, DOLOMITE. Clay with 40% sill, 5-1b% very fine sand. Arenaceous fraction of

moderately sorted, angular grains, Colour 3Y6/L. orange to orange-brown mottling, Nodules and

patches of greenish-white very line grained sandy dolomite, weathering ax 2 cm granules al top and

bottom, Black stained,

0.78. Alternating SAND and CLAY. Sand beds range from silt to very fine sand, colour 5V8/2. Clay is

sandy. Individual beds 2-27 cm. Lower beds have shatper contacts and are laminated, very finely cross-

bedded and lensing. Grange brown mottling and gypsum) iy upper part of section. Miner tedulur dolo-

mile,

0.80, SILTY CLAY, Silt to very fine sand 10-20% svbangular frosted grains. Clay bas greasy lustre

and SY6/2 colour. Upper contact very sharp, Ast.

1.00. SAND and DOLOMITE, Sandy clay and clayey sand with dolomite nodules 2-30 om thick at top

and bottam, Sand is subanyular to subrounded. moderutely well sarted, with frosted grins: Weathers

to hard Jight brown sandstone, base cemented with gypsum, Sandy clay is S¥5/Z, clayey sand S¥7/3,

MmotUed with yellow green patches, and black pateles at base (NI-NS5). Very sandy parts are INYRG/2-

SYRS/2 ut buse.

Dorel is very fine grained with botryoidal upper surface, in way, lensing beds. Greenish upper sur

aces,

0.30. SAND, As above,

0.33. CLAY. Mottled clay and very fine sand, finely Juminated beds which ate irregular and lenticelar,

and have yellow green (10¥R6/6} clay patches and, cross-cutting brown (10YR8/6) patches with black

centres. Sand (5¥7/2). Basal 2 cm fine white sand, Gradational upper contact. Contains dark brown to

yellow-orange irregular silcrete and ironstone nedules.

0.02-0.03. CLAY. As above. Sharp contacts.

0.23. SAND. Very fine, yellow green clay lamellae. Obscure medium cross bedding, straight foresets.

Stained greenish black. Sharp upper contacts,

0.25. CLAY. As hieher in the sequence. Upper S-[0 cm dolomitic, yellow-sreen and brown mottled.

0.10 SAND, As above. Obscurely cross-bedded, selenite cemeni al base.

1.30. CLAY. Silty, Very finely laminated, with very small scale cross-hed sets. Colour SY6/0,5, aces.

sional black and brown patches, Upper contact sharp, flat,

0.70. CLAY. Slightly silty, hard, with sub-conchoidal fracture and ereasy Justre. Selenite on contact with

overlying unit. Stight greenish brown dint, otherwise N4.

Section 138, south of the nerthern track, crossing Lake Tarkarnoloo (200 m sowh of section J3A).

Constructed from a series of breakaway slopes, and a scoured channel In the centre of the lake,

Beginning from the top of the black clay, as al the base of the previous section, the sequence is us

follows (from gally sequence};

3.5. CLAY. Dark vrey elay with irregular shiny surfaced fractures, becoming IHehter coloured towards

base. Crops out poorly, forming low angle vegetated slopes, Upper contact sharp, flat. Lewer contact

moderately sharp to disturbed.

2.1. SAND, Very fine to fine gruined, black stained. Thin S cm beds and fine laminate, Brown silcrete

nodules throughout, Thin horizon of DOLOMITE nodules ac top, Abour 20 cm above the base are 10

cm of laminated silty limestone with low amplitude symmetrical ripples. mud cracks, and 1-3 mm

tong tubules of organic origin, Sharp basal contact with -, -

0,25. CLAY, Ax al lop of sequence (13B).

On a small nol isolated near the edge of the fake, the sequence continues os follows, with some overlap;

0.55. DOLOMITE, Sandy nodular whire dolornite. Impregnated with xypsum and calcite, plus tabular

black manganese concretions.

0.98 SAND, Fine grained, well sorted, laminated and silty at base. Cemented by gypsum and calcite in

part. Numerous curbonate nodules cahibiling concentric structure. Some have vertical tubular disposi-

tian with am intemal structure snggestive of shrinkage (c.f, silorete nodules}, Hlack and brown stain at

base. with orange brown and yellow Sreen patch stain. This, and the previous unit are equivalent to rhe

2.) m of sand described above, with its hed of dolomite nodiiles.

2.5, CLAYEY SILT. Alternating hard clay and clayey sift in very fine Jamellae (varve-like), Silty grey

clay lenses. Yellow green patches with waxy lustre. Clay dominant at base. N35, IDYS/4 to 4'6. Con-

tact with underlying unit sharp, undulating: sume contac! as at base of 2.1 m sand described in paily

section abave.

0.5, CLAY. Hard, areasy lustre, irregular Tracture. N3 at top grading down to SY¥4/I,

This clay crops out across the bed of the Jake jo its centre, where 4 scour next to a salt spring exhibits

a further 2.5 moof massive hand ercy clay,

158

R. A. CALLEN & R. H- TEDFORD

NAMBA FORMATION SUPPLEMENTARY SECTION

SADM Wooltana 1 Stratigraphic Bore. Section Ll, Fig. 3

°EURINILLA FORMATION

0.0 1 5.2 m No core

3.2

0.9 COBBLE CONGLOMERATE: Cemented with réd-brown carbonate, Mica schist, gneiss,

quarizite. Sindy (medium grained), with subrounded to well rounded, erains,

— ?Disconformity —

WILLAWORTINA FORMATION AND NAMBA FORMATION {intertonguing})

Units

7.2

iA)

12,50

28.65

33.00

38.6

43,05

48.00

53,15

67.65

NAMBA

Units

74,35

1.1 DOLOMITE. Sandy, clayey at base. SYR6/4 to N6. Sharp upper contuct, gradutional

lower contact,

3.8 CLAY. Slightly sandy, with dolomite nodules. Clay 7¥7/2 with red-brown vertical pipe

structure, Sund subangulur to subrounded. Some gypsum patches im lower part, Lower con-

tact gradational, 50 cm core missing,

0.65 SAND, Fine sand, very poorly sorted, subangular to subrounded, Thin slntomite beds.

Dolomite nodules, Clay at base.

11.65 to 12.50 Na core

16.15 CLAY erading to SAND at base. Numerous dolomite nodules and sume beds,

Extrentely. poorly sorted mediim sand to coarse sill, Some gypsum patches at top. Green and

red-brown pipe structure, Sand subangular to well rounded, Sharp basal contact. 40 cm core

missing in sand interval,

4.35 CLAY grading to SAND at base, As above. Mica in basal sand. 2.5¥R/4-6, 5Y6/)

moriles. Sharp wavy basal contact.

$.6 CLAY, and DOLOMITE erading to SAND at base, As above, but dolomite beds in upper

clayey part, Dolomite in 20%. clay, Medium sill, extremely poorly-sorted. Sand subangular

to rounded. Gradational wavy lower contact.

4.45 CLAY, grading to SAND at base. As above. Dolomite beds (brown) and nodules

throughout. Sand patches al top. Lower contact eradatianal,

4.95 CLAY grading to DOLOMTTE then SAND at base, As above. Mouted red-brown, green

and yellow grey, Very poorly sorted. Minor core loss. Lower contact gradational,

5.18 SANDY CLAY grading to SAND at base, As above, Dolomite beds and nodules

throughout, Sand laminated, micaceous. Intraformational clay and carbonate at hase, Sand

reaches medium grain size. Subangular to rounded, Disturbed irregutar basal contact.

14,50 CLAY cf. NAMBA FORMATION, Patchy sand and carbonate near top, Reticulate nét-

work of carbonate “veins®. Lower part with limonite nodules, irregular shiny blacke-stained

fractured clay. Obscurely laminated in middle part. Sand very fine, angular to subrounded.

SY6/1, 2SYR6/2-8, 1OYRB/2, 2.5YR5/6, SY7/1.5, SY6/i, SY¥4/5, 2-3 m core missing,

mainly in upper part.

.7 SANDY CLAY. Micaceous finely laminated silt becoming pebbly at 72 m, reverting to

clay at base. Extremely poorly sorted, Granite, quartzite, shale, quariz and eneiss pebbles,

Very angular fo subrounded. Clay intraformationally brecciated and burrowed in lower part.

Mincr carbonate, c.f, WILLAWORTINA FORMATION, Gradational lower contact.

FORMATION

4.9 Alternating DOLOMITE atid CLAY, Dolomite (5-10 cm thick), oolitic white, aphanitic.

with charophytes, gstracodes, molluscs and unidentified calcareous %plant fossils. Numerous

burrows in clay beds (all about 1 m thick). Micuceous silt in part. Bioturbated, Laminated

at base, Sharp basal contact.

7.9 CLAY, minor DOLOMITE. Upper purt similar to above, laminated and burrowed, Clay

GYO/T+ ZY RAG, becoming sandy at base, With rounded clay clasts, Basal contact grada-

tional,

1.75 SANDY CLAY. Calcareous. SY5/1 to 5¥6/1. Sharp basal contact.

0.10 CLAY. Sharp contacts.

2.00 Interbedded DOLOMITE and CLAY. Dolomite as above. Very finely interlaminated with

brittle, swelling clay, Shrinkage ¢racks common. Burrowed, Contacts on carbonate bedé sharp

and wary to disturbed irregular, 7¥571, 5¥6/1, N3,5, 5¥4/2.

2.55 MARL and GYPSUM. Alternating thin selenite atid calcareous clay, Sharp contacts of

gypsum. Gradational lower contact. Black to dark olive.

4.60 CLAY grading to DOLOMITE and MARL at base. Numerous gypsum nodules, 4¥5/1,

FY8/2. Mottled SYR6/7. Minor gypsum lAminac, Contacts wavy eradational to disturbed

irteguiar. Intraformational brecciated,

3,35 CALCAREOUS CLAY. Trregular shiny Fractures, oxidized red-brown patches. Swelling,

very porous. Subaqueous shrinkage cracks,

$15 CLAY. Sandy in centre, with selenite veins infilling slickensided joints. Sand laminated.

Clay with irregular shiny surfaced fractures as above, 10-15% very fine sand to silt. Alternat-

ing colour pattern—oxidized red-brown clay passes down io 3¥4/2 10 6Y7/1 clay, has sharp

upper surfaces. Basal contact regular, disturbed.

7.L CLAY. As above. Busal thin white dolomite. Churmed structure suggests biaturbation,

TAS CLAY grading to SIT.T in lowe half. Silt well laminated, micaceous, very small scale

125.45

129.00

130,80

138.00

140,40

Unit 3

147.30

163,40

165,20

166.80

170.80

175.30

183.7

187.3

196.10

Unit 2

197.0

241.0

2273

236.0

CAINOZOIC ROCK UNITS 139

cross-laminated, Grains yery angular to sub-rounded, Silt moderately sorted. Lower contact

moderately sharp. :

4.55 Alternating SAND, CLAY. Sand very fine, micaceons, small scale cross bedded,

laminated, Lower contact moderately sharp,

1.80 As above. Lower contact irregular, wavy,

$5.00 SAND, interbedded CLAY, Sand fine to medium, well sorted, small to mediutn scale

eross-bedded_. Micaceous and with clay balls. Grains very angular to subrounded. Sharp wavy

contacts.

2.60 SAND, As above, Upward fining, Some clay at lop, burrowed. Poorly sorted, average

very fine grained. Wavy moderately sharp lower contact,

6.70 Aftemating SAND, SILT, CLAY, CARBONATE. Sand as above. Carbonate nodular.

Clay dark grey. Bioturbated and intraformationally brecciated. Mainly fining upward

sequences, Sand dominant. Contacts irregular, disturbed.

26.140 CLAY grading to SAND in lower 1.5. m, Manganese nodules above sand. Minor sand

bed at 157 m. Cloy N45, numerous imegular shiny fractures. Sandy, averaging very fine silt,

very poorly sorted. Sand and very fine silt, very poorly sorted. Sand very fine, burrowed and

laminated. Very angular to angular grains. Irregularly disturbed lower contact.

1.8) SILT. 5Y6.5/0.25, orange brown moltles, obscure lamination, bioturbated. Sharp wavy

fower contact. Grades io very fine sand at base,

3.4 SAND, Fining upwards from fine sand to clay. Thin clay bed with sharp contacts near

tup. Medium scale cross-bedded. 6Y6/1 to S¥7/2. Sand poorly sorted. Moderately sharp

wavy lower contact,

4.00 CLAY grading to SAND at base. Upper | m sandy, lowee very fine obscurely laminated

sand, Clay with irregular fractures, SY4/1. Sand beds have gradstional contacts,

4.4) AS ABOVE. Minor sand at 172.2 m, Basal very fine sand, moderately sorted, obscurely

small scale cross laminated, burrowed, Gypsum nodules. Clay as above with fractures and

orange [6 red-brown moltles. Sund very angular to angular.

74 Silt. Laminated fine, very poorly sorted silt, xypsiim nodules at top. very small scale

cross-bedded, some burrows. 3m missing im central part. Sharp wavy lower contact.

4.6 SANDY CLAY, gradmgz to SAND at base, Cycle as above, Some calcareous zanes, SY¥5/1,

SY3/4, Sandy at (85 ny, Basal sand fine graincd, sub-rounded. Moderately sorted, Lower cou-

tact moderately sharp.

&.8 SANDY CLAY, SAND at base, Upper 3/4:N2 to SGY7/0.5 clay as above with fractures

and sandy patches. Dolomite nodules at contact with sand. Sand very fine to fie, very poorly

sorted, SY6/! to N&, Burrowed near top sand, rest Wloturbated, obscurely cross-stratified,

0.9 SILT, Minor alternating clay and sand, sharp contacts. Silt laminated, Lower contact

shafp and wavy.

14.0 Alternating DOLOMITE, CLAY, SILT and SAND, Complex inter-relationship between

clay-dolomite cycles and sand and clay beds c.f. above. Contacts variahle. Dolomite intra-

formationally brecciated, Sut laminated. Bioturbated and burrowed horizons. Clays with

iffeeular fractures and orange mottles. Beds 40 cm—1-5 m thick.

163 Alternating LIMESTONE, DOLOMITE and CLAY. Consists of 1-2 m carbonate beds

grading up to clay via, disturbed zone, Clay beds burrowed or biolurhated, |OYS/2 to

IGYR7/2. Base of dolomite beds sharp und wavy. Dolomite aphanitic, white laminated, with

oolitic zone. Ostracodes common, algal mats present. Zone 215-217 m of very narrow clay

filled cracks, Irregular shiny fracttres dominute in clay near base, otherwise absent, At top of

this sequence is 5 em genethite-limonite crust.

8&7 CLAY. Calcareous, intraformationslly brecciated with numerous white carbonate specks,

Irregular shiny surfaced fractures, N1 to olive green, Quartz rare, very fine to fine, angular,

Lower contact rradational.

8.45 CLAY, Fissile pyritic carbonaceous very finely laminated clay with sift parting. Fine

laminac of N1 to dark olive or SY2/1 colour’ Numerous plant stem and leaf impressions or

fruiting bodies, fish spines and scales, ostracodey (often in pure layers), gastropod prato-

conchs, spores and pollen. Burrows (pyrite filled) and bedding plane traces. Numerous pyrite-

marcasite nodules. Some subaqucous shrinkage cracks.

WILLAWORTINA FORMATION TYPE SECTION

Werstern Nuclear WC2 Bore. Section 10, Fig. 3

*MCOONARBINE FORMATION and JEUIRINILLA FORMATION

0.00 to 7.05 Cuttings only, SANDY PEBBLES, SAND. Micaceous. calcareous, impregnated with gyp-

sunt (except sand). Angular to very angular, 2. 5¥R4/8, 2.5YRI/6,

— Disconformity —

WILLAWORTINA FORMATION—MEMBRR 3

7A5

7.89

$54

984

0.44 No recovery,

1.65 SAND as below, ]-39% muscovite.

0.30 No recovery.

3.95 SAND grading to SILT. Sand very poorly sorted and very fine erained, silt coarse, pebbly

160

R. A. CALLEN & R. H. TEDFORD

and micaceous. Thin pebble bed grading up to sand at base, Porous zones impregnated with

gypsum,

0.25 No recovery. .

1.21 PEBBLY CLAY SILT, poorly sorted, with basal pebble bed, impregnated with gypsum.

1.50 No recovery.

0.24 PEBBLES, subrounded to rounded, Coarse feldspathic gneiss, fine biotile quartz feld-

spar gneiss, purple stained coarse feldspar,

0.68 No recovery.

0.13 SAND, pebbly, coarse grained, Feldspars, muscavite, biotite

1.98 No recovery—a few abraded pebbles.

0.24 SAND, fine grained and micaceous.

0.22 No. recovery,

0,38 GRANULES grading up to PEBBLES.

1.45 No recovery.

0.28 GRANULES to SAND, medium grained. Larger grains subrounded. Mica 5%, pink

potash feldspar 15-20%.

{).64 No recovery.

0.33 CLAYEY SAND, very fine grained and micaceous, sharp wavy content with overlying 20

cm of COBBLES. 5-10% muscovite, pink potash feldspar. Yellowish grey (SY7/2Z).

1.49 No recovery.

0.11 SAND, grading up to GRANULES. Sand medium grained with dull, pitted or shiny sur-

faced, angular to subangular grains. 15-20% feldspar (mostly pink potash variety). Musco-

vite and biotite flakes in the quartz.

G.80 No recovery.

0.20 SAND, pebbly, micaceous and overall fine grained.

0.72 No recovery.

0.14 COBBLES, sandy.

0.47 No recovery,

0.17 Pebbly micaceous coarse sand.

1.01) No recovery:

0.27 SAND, grading over short interval to COBBLES at top. Sand is poorly sorted and

micaccous, medium grained.

1.61 No recovery.

0.27 COBBLES, passing over short interval to medium micaceous SAND,

0,72 No recovery:

0.53 SAND, grading to COBBLES in upper half, ‘Sand micaceous and poorly sorted, medium

grained, Clasts pink quartzite with mitaceous hematite, purple fine quartzite, vein quartz.

large potash feldspar pebbles.

1.00 No recovery.

Q.26SANDY PEBBLES, micaceaus.

0.58 No recovery.

0.30 SAND, coarse and poorly sorted, micaceous.

0.30 No recovery,

rey PEBBLY SAND, very poorly sorted, micaceous, coarse grained. Dusky yellowish grey

5Y¥6/2).

0.57 No recovery.

0.24 CLAY, sandy, micaceous,

0.62 No recovery.

0.13 PEBBLY SAND, fine grained, moderate reddish brown (2.5YR5/6 to 4/6),

2.00 No recovery. f

0.30 CORBLES and PEBBLES overlain by fine grained CLAYEY SAND.

0.20 No recovery,

9.56 GRANULE bed, thin, overlain by thick SANDY COBBLY PEBBLE bed (white vein

quartz, pink potash feldspar, pink ferruginous quartzite, coarse siliceous gneiss or granite, dark

grey Shale, weathered fine gneiss).

0.77 No recovery.

0,20 SAND as before, with clayey sandy cobble bed at top.

0.71 No recovery.

2.46 SAND, coarse at base, grading up to extremely poorly sorted fine sand. Vertically

oriented reddish brown pipes.

().29 No recovery,

0.53 PEBBLY SAND, sand medium grained.

0.23 No recovery.

2.77 Fine SAND as before, grading up te pebb)y medium grained sand in upper | /3.

0.18 No récovery-.

1.47 SAND, poorly sorted and very fine grained, micacteous, with scattered pramiles, two thin

pebble beds at base.

0.63 No recovery.

0.35 PEBBLES, passing over short interval ta SAND, clayey, micaceons and fine grained.

poorly sorted,

CAINOZOIC ROCK UNITS 161

50,05 1.64 No recovery. _

51.69 a CLAYEY SAND, coarse and very poorly sorted, grading to boulders. Metaquartzite

clasts.

52.03 1.23 No recovery.

53.26 0.29 SAND, micaceous and yery fine, light yellow-brown (7YR5/6). Very poorly sorted.

53.55 125 No recovery.

54.80 0.50 SAND, very micaceous, very fine grained. Pebble bed in centre (granite, banded pink

and white quartzite).

55.30 0.58 No recovery.

55.80 0.63 SAND, fine, grading up to fine grained with scattered granules.

56.43 0.30 No recovery.

36,73 0.44 SAND, as above. Feldspathic quartzite pebbles.

57.17 0.48 No recovery.

57.65 0.40 SAND, as above, pebble cobble bed in centre.

58.08 0.49 No recovery,

58,54 0.84 CLAY, and pebbly SAND, coars2 grained. Grades rapidly to micaceaus CLAYEY

SAND, fine grained, in upper 30 cm.

59.38 0.69 No recovery.

60,07 0.56 SAND, as below, fining to fine grain size at top,

MEMBER 2

60.63 0.66 No recovery.

61.29 0.54 SAND, as below, no granules.

61.83 1.43 No recovery.

63.26 0,99 SAND, slightly clayey, medium grained, with granules. Coarsening upwards.

64.25 1.50 No recovery.

65.75 Ps CORRES: massive pink granite, very fine dark quartzite, pink feldspar with ?horn-

ende,

65.98 1,30 No recovery.

67.28 0.10 SAND, medium micaceous and clayey.

67.38 0.19 No recovery,

67.57 0,34 GRANULES, grading to CLAY-SILT.

67.91 0,84 CLAYBY SAND, coarse, pebbic interbeds.

68.75 0,05 No recovery.

68.80 0.47 SAND, coarse and poorly sorted, pebbly, Pebbles of quartz and gneiss.

69.27 0.14 No recovery.

69.41 0.41 SANDY CLAY, as before grading up to COBBLY SAND.

69.82 0.12 No recovery,

69.94 0.18 SANDY CLAY, as before. Micaceous.

70.12 0.25 No recovery.

70.95 0.85 Silty clay grading up to cobbles,

71.82 0.33 No tecovery.

72.15 0.27 SAND; medium, grading up to CLAY.

72.42 0.13 No recovery.

72.55 4.07 SILT, extremely poorly sorted, medium size.

76.62 0.11 No recovery.

76.73 1.40 SAND, very fine, pebbly, grading to SANDY SILT CLAY.

78.13 0.95 No recovery,

79.08 0.85 GRANULES (lower 20 cm) grading up over short interval te SAND, clayey, medium

grained, very poorly sorted,

79.93 0,17 No recovery.

80.10 1.68 SAND, very fine, grading to very coarse at top. 1% miuscovite and biotite, 10-15%

potash icldspar. Grains very angular to. subangular and dull, Small grains shiny and faceted.

Extremely poor sorting,

81.78 0,15 No recovery.

81,93 0,25 SAND, fine grained, very poorly sorted.

82.18 0,16 No recovery.

82.34 ee GRANULES, basal bed, grading over shoft interval to very poorly sorted CLAY, very

sandy

MEMBER 1

83.18 0.06 No recovery.

83.24 0.76 CLAY, lower 10 cm sharp contact with SAND, coarse grained to granule sized. CLAY

thin bed at top. Extremely poorly sorted,

84.00 0.37 No recovery..

84.10 0.10 SAND, fine grained, clayey.

84.47 1,37 No recovery.

85.84 1.50 SILTY CLAY. Very poorly sorted, with thin coarse sand beds.

87.34 2,03 No recovery.

89.37 5.74 SILT, SILTY CLAY, Extremely poorly sorted coarse silt, silty clay micdveous. Thin

coarse grained sand bed (91.2 m), above which the sediment coarsens from clay to very fine

gtained sand.

(62

95.11

95,36

97.37

97,63

101.42

102.40

104.80

105.90

110,80

121.20

21,90

123.35

126.45

R. A, CALLEN & R. H. TENFORD

0.45 No recovery. ;

1.8) SAND, grading to CLAY a1 top, Sand very coarse, clay silty and micaceous with a thin

eranule bed near the top. Yellowish grey (SY8/1)-.

0.26 No recovery. , .

3.79 SANDY CLAY, SAND. Sandy clay has medium grained sand fraction, very poorly

sorted, 2rades to clayey coarse sand, with very angular lo subangular pitted to shiny grains,

Feldspar is common. The base of this interval is taken as the base of the WILLAWORTINA

FORMATION.

1.08 No recovery.

NAMBA FORMATION ;

2.30 CLAY. 15-20% subanvular to subrounded sand, minor mica. Sand patchy near base,

with irregular shiny-surfaced fractures (skew-planes), SY6.5/2 to 4¥5/1. Basal contact sharp.

30 cm Core missing neur base.

1.40 CLAY. As sbove. 10% sand, no mica, nodular and swelling with well developed frac-

tures. Alunite mottles at base, 20 cm core missing in centre of interval, 1Y4/1, to 6YR6/1 at

base, Sharp basal contact. :

4.90 CLAY, grading to SILT ip basal 60 cm. Intraformationally brecclated and burrowed (at

109 m} with some laminated intervals. Well developed alunite streaks, which decrease in abun-

danee with depth, being absent al the base. 6YR6/1-6/2 grading to N8 at base, 70 cm core

missing al vatious intervals, mainly near top. Basal contact irregular, disturbed.

10,50 CLAY grading to SILT at 144.4 m and SAND at 119.4 m. Clay intraformationally

breceduted, 15-25% very angular to subrounded sane. Silt micaceous, Sand pnicaceous, cross

bedded in 30 cm sels, and Jaminated, fine grained, well sorted, Grains angular to subrounded,

Basat contact gradational, Weak alunite horizon 50 cn below top of unit, absent at 115 m.

Colour 5¥YR5/1 above alunite, N8 below, 2,1 m core missing in silt and sand beds. Wavy in-

diatinet luwer contact,

0.70 SILT. L

\.45 SAND. Micaceous, Jaminaled, obscurely croas-bedded_ Fine grained and moderately well

sorted. Shurp lower contact, 70 em core missing in centre of unit.

3,10 SAND. Minor clay at top, fine grained micaceous sand in centre, lower half grading to

very coarse sand iat base, Subangular to subrounded, large grains highly polished composites.

Mostly no core, there being 50 cm recovered. Basal contact sharp,

9.45 SILT and CLAY (below 129 m). Clay, nodular, dark brown, sill greenish white. Sandy

patches, Sand grains often show crystal faces—bipyramids. Indistinct contacts observed at

129,25, 129.4, 129,6 associated with weak alunite horizon, Colour 1OYR6/2-5Y8/1 in this

zone. Below 131,80 irregular shiny surfaced fractures and some alunite specks, Colour

SY5/1-4, SY3/1. 17% silt. Grains very angular io angular. Much core missing throughout.

recovery 40%.

WILLAWORTINA FORMATION SUPPLEMENTARY OUTCROP SECTION

—'Discentormity —

“Wertaloona” Homestead Arca. Section L, Fig. 3

WILLAWORTINA FORMATION

Unit $,

Unit 8,

37,4

Unit 7-

37.1

Unit 4.

107.0

Unit 4.

Unit 3.

Unit 2.

Unit ft.

140.1

37,9 COBBIFS, Brown quartzite cobbles in a matrix as for unit 4. Basal bed of almost 100%

erey-blue limestone cobbles. Rare red sandstone, quartz and yellow-brown silicified. carbonate

cobbles in float. Exposure poor, top nat exposed.

49.2 SANDY CLAY, red brown,

20.8 COBBLE to BOULDER beds. Matrix as for unit 6, cemented with secondary white car-

bonste which may be powdery and soft, or hard vughy and crystalline. Cobbles of brown

quartzite with 20% blue-grey fimestone (resembling Cambrian limestones). Rare very large

boulders of grey massive microcrystalline quartzitic silerete with large milky quartz pebbles.

(3.0 (approx.} CLAY SAND. Red brown very poorly sorted and calcuréous,

2.0 (approx.) COBBLES. Brown quarizite cobbles scattered through matrix as for unit 4-

Lenses out along strike.

7.0 (approx.) CLAYEY SILT SAND. As for unit 2.

1.0 (approx.) PEBBLY COBBLES, As for unit 1, more matnx, thin and lensing along strike.

a8 {approx.) CLAYEY SILT SAND. Red brown, with a calcareous matrix, sametimes thinly

aminated,

4.0 (approx.) COBBLES. Brown quartzite pebbles and cobbles in calcareous red-brown silty

sand, lenses of calcareous medium sand at base. The sandstone fills channels, which have

groove casts on the base. Cementation is weak, and pebbles weather aut readily with thin cul-

careaus crusts, Proportion of matrix tow. The unit cuts into deep red brown clayey silt,

probably Namba Formation. Althaugh the contact here is sharp, there may be an inter-

tonguitig relationship along strike. The unit grades laterally to the south into pebbly clayey

satidstone,

The following part of the section is poorly exposed, and is yet to be fully described:

— Disconformity —

2NAMBA FORMATION

(2,0 SAND. Very fine greenish sand grading wp inte silty grey-green clay with gypsum patches.

CAINOZOIC ROCK UNITS 163

7.2 CALCAREOUS SANDSTONE. Very angular sand with soft crystalline carbonate cement Contains

pebules of very angular sandstone, carbonate, rounded brown quartzite, polished milky quartz, chert

ranules.

51 2. CLAY. Dark gréeen-grey clay with greenish-yellow-stained patches, slightly sandy, Thin white nodu-

far dolomicrite is preseml near the base, aid may be a facies variant of the previously described nit,

3.2 SAND, Reddish to greenish silty fine to medinm sand.

$2 CALCAREOUS SANDSTONE. Essentially a sandy limestone with aboui equal quantities of

medium grairied angular sand and lime. Weathers grey, With a sculptured ‘rough surface.

144.7 CLAY, Grey green to olive, greasy irregular fracture, sandy and silty. Minor dark olive to grey

clay. Mottled with red-brown iron oxides. The interval 430-350 m (measured from top of the unit 9

in the Willawortina Formation section) is very poorly exposed and deeply weathered.

Near the top of these beds in the northern part of the ares, is a thin white dolomicrite bed,

— Discanformity —

YEYRE FORMATION _

2.0 SANDSTONE. Massive calcareous medium grained sandstone, partly silicified, and capped by rem-

nant silcrete, dipping with the section. ‘

2,0 CONGLOMERATE, Granule to pebble-sized polished white guartz, grey chert, ironstonc. Pebble to

cobble-sized angular Middle Cambrian sandstone. All in medium well-rounded sand matrix, cemented by

calcium carbonate. Pebbles are patchily distributed, and the whole crops cut as a Jow ridge wath caver-

nous weathering and of brownish grey colour, Medium scale cross-bedding is prominent. The unit has

an apeplar unconformable relationship with the underlying Middle Cambrian red-beds, though dips arc

aimilar.

Appendix Ti

YOUNGER CAINOZOIC UNITS

FIG, LS SECTIONS 2-9

(See Fig. 1 for locations, and maim exe for access and photo points)

SUPPLEMENTARY SECTION, COONARBINE FORMATION

SECTION 2

COONARBINE FORMATION

1.7 SANDY SILT. with basal pebble bed. Sand dark brown (5YR3/5}, Size varies from silt to very fine

sand, moderately poorly sorted. No bedding planes Visible. Columnar structure well developed. Basal

clasis may be small cobble size, and are of metamorphic rocks and quarz,

— Disconformilty —

EUXINILLA FORMATION

2.2-2.5 CLAYEY SILT-SAND. Very poorly sorted. with irregular-shaped frosted or pitted grains, Con-

lains pebble lenses (though not in the figured section) and large irregular aphanitic greenish white

sandy carbonate limps. The latter are probably derived from the upper carbonate in Wooltana 1 bore

{section 1, Fig. 3). At top of O.5-1 cm diameter branching vertically oriented cylindroids of pinkith

“chalky” textured carbonate, representing a fossil soil horizon.

— Disconformity — .

0.2 CALCAREOUS SAND. Pebbly sand (coarse grained), solidly cemented by pinkish buff (SYR7/2)

carbonate, Colour derived mainly from orange-stained quartz grains. Laminated and thin bedded, Beds

dip, suggesting cross-bedding is present (outcrops seen in plan only, in creck bed), :

Possibly represents Willawortina Formation, or unnamed conglomerate equivalents of Millyera Forms-

bon.

SUPPLEMENTARY SECTION, COONARBINE FORMATION, EURTNILLA FORMATION

SECTION 3

Location, Curnamona Siccus map sheet, Air pheto ref.:; S. Aust. Dept, Lands Svy, 361, run.2, photo no,

4442. The section is situated on the northwestern bank of the Pasmore River, close to the point where :t

debouches in to Lake Frome,

RECENT

0.00=1.20 Mobile bright red-hrown dune sand, sharp erosional contact with underlying units.

— Disconformity —

COONARBINE FORMATION

1.90-3.50 SAND. Yellow-brown, with large scale dune-type cross-bedding, Sharp erosional basal con-

tact, A lag of pebbles (eroded from the Eurinilla Formation) is al the base,

Numerous broken Mature snail shells are present in the upper part of the unit. Aboriginal artifacts, cal

sified iree roots, emu shell, and vertebrate bones occur in the uppermost. level (or possibly on the

upper surface in the case of the artifacts and emu shells).

Strongly developed columnar structute is present (resulting from soil processes).

— Disconformity —

EURINILLA FORMATION

1.80 PEBBLY CLAY-SILT and SAND, Sand at base, medium-grained, yellow brown, numerous

pebbles and care flat cobbles, cemented by gypsum. Pebbles are milky and clear quartz, and very angu-

lor frugments of calcite-cemented conglomerate, overlain by bright red-brown silty clay

Unnamed Conglomerate (7Millyera Formation equivalent)

fh.15 CONGLOMERATE, Thin, calcite cemented. Pebbles weather out without adhering crust. Pebbles

164 R. A, CALLEN & R. H. TEDFORD

as for overtying unit plus ?7Namitha Formation dolaomictite. and brown carbonate nodules from Willawor-

tina Formation. Carbonate penetrates into top of underlying bed,

— Discontormity —

WILLAWORTINA FORMATION

2.05 SILTY CLAY. Sandy, preenish-brown with red-brown motiles. hard. Patches of gypsum nedules.

Partly calcified at top. Blocky columnar structure visible (resulting From soil processes). Upper contact

sharp, undulating. ;

MILLYERA FORMATION TYPE SECTION

SECTION 4

030 SAND. Reworked from older unit into base of dunes,

0.70 SAND. Coarse grained, with many gypsum grains and anomalous pebble sized angular quartz

(milky), Powdery hummocky gypsum often developed at top ¢sojl profile),

— Disconformity —

BRURINILLA FORMATION

1.10 CLAYEY SAND. Very fine-grained, sub-rounded to rounded. good sphericity, moderate sorting.

Numerous charophyté oogenta. Muny greenish, yellow and brown grains, Colour S5YR5/6, Capped by

£ypsum crust, of gypsum nodules in clayey sand (groundwater deposit),

— Disconformity —

MILLYERA FORMATION

Unit7, 0.50 CLAY. Soelft, conchoidal fracture. Contact with overlying Eurinilla Formation sharp and

flut. Very dark yellowish brown. The oxidized crumbly appearance and shiny surfaces (cutans)

on cnimbs suggest soil processes have operaled, and indicate a disconformity between Millyera

; and Evrinilla Formations, _ .

Unité, 0,40 SAND. Very fine to medium erained. Grains subangular to rounded and frosted. Charo-

, phyte oogonia .5%. Grades by alternation, 1a... |

Unit 5, 0.50 CLAYEY SAND. Sand fraction well-sorted, with subangular to angular rounded grains,

sharp fiat upper contact, Greenish yellow (1077/2).

Units, O70 CLAYEY SAND, Interbedded thin clay and very fine to fine clayey sand 0.25-0.50 mm

thick. The sand iv very well sorted, with subrourded to rounded high sphericity frosted grains.

Darker oxidized clay present, Yellowish grey (7¥7/2). Lower contact gradational,

Unlt3. 0.30 CLAY as for unt 1.

Unit2, 0.40 LIMESTONE and CLAY, Near the top of the sequence each cluy Jamina grades up to

charophyte stem-mould limestone (up to LE cm thick), These limestone beds harden on weather-

ing, producing sheets and slabs with » metallic ring when struck. Intervening lamellae are 0.5—

2 om thick. Some of ihe Charales tubules ar¢ oriented and small turreted gastropods. aif.

“coxiella” ure present (henceforth referred to as “Cextella). An oxidized zone exists beneath

the limestone. The limestones form.a distinctive marker horizon 20-30 cm thick. Contact with

unit | and unit 3 are gradational by alternation.

Unit!, 0.10 CLAY, brittle, soft, waxy lustre. Distinctly laminated and thin bedded (1-5 cm), each

lamina grades up to # thin fine silt layer wilh charophyte oogonia and Ostracoda, White car-

bonate granules occur near the base of the sequence, Scattered medium polished or frasicd

quarlz grains, sometimes up to 40% of the rock, occasionally forming sand lenses. Vellowish

grey (5¥6/2—+lay, lighter for sand), Base fot exposed.

SUPPLEMENTARY SECTION, COONARBINE FORMATION, EURTNILLA FORMATION, MILL-

YERA FORMATION

SECTION §

0.00-3.50 SAND, Red brown sund of modern dunes reworked from Coonarbine Formation.

COONARBINE FORMATION

1.00 SAND, Light brown. Numerous vertically oriented small cylindroids of soft white carbonate, of

soil profile. Emu shell, aboriginul artifacts and rare mature snail shells Gocur in uppermost level, Forme

longitudmal dunes.

— Disconformity —

EDRINILLA FORMATION 4

Unin2. 4.060 SAND, fine to medium prained, with subungular rough of pitted grains, poorly sorted.

Stratificatian absent. Grades to unit 1 over short distance. Light brown (SYR4/'7)-

Unitt. 4.00 SAND, medium grained, brown (SYR6/H), lighter coloured beds alternate near base,

Cross-hedded, scts 40 cm, lensing, gently curved coarse and fine laminae, sharp ereded wpper

contacts, assymptotic bottomsets. Laminae &S-l cm, by variation in clay content. Sets are

gently inclined toward Lake Millyers. Numerous charophyte oogonia.

Lightly cemented with clear or white finely crystalline carbonate. Pinkish irtegulay nadules,

weathering as brown lumps and slabs on surface. Carbonate gives white cast to This part of the

section, and causes slight benching, Partly cemented with massive gypsum in the basal layers,

— Disconformity —

MILLYERA FORMATION

Unit, 7,60-2,450 CLAY. Very hard, shiny irregular fractures, coated with black iron oxide and While

carbonate al top (soil horizon). Impregnated with vertically oriented gypsum masses. in 5S—

10 cm columns (fossif groundwater horizon) at top. No silt content. Colour 10¥6/2, Similar

fo Willawortina Formation. Upper contact sharp, flat. Grades down to light green soft clay tn-

lérbedded with very fine grained white sani) nch in charophyte oogonia Intertongues with

unit 4,

CAINGZOIC ROCK UNITS 165

Unit4. 4.00-5.00 SAND. Silt to very fine grained sand, with coarse lenses. Numérous thm ,5-5 cm

clay beds and lameline near top, which are crowded with algal tubules (charephytes}, Some

Tare massive charophyte crusts consisting entirely of strap-like algal forms with mutherous

large oogonia, Charophyte oogonia commion in upper sands. Clay pellet layers common. Sand

grains are subrounded to well-rounded smooth or frosted, with moderate sorting, Small scale

cross-laminated sets, 1D cm thick, with curved laminae. ;

Upper surface may be cemented with gypsum of a fossil groundwater horizon.

Unit 3, 0,00-0.93 SAND. Very fine grained, well sorted. Colour 5YR6/8. Impregnated with massive

Eypsuns and disc-shaped crystals of gypsum, Grades by alternation of 1-4 cm thick beds into

overlying unit, in which rt forms a lense, Contacts between lamellae are wavy, lenticular, and

Tippled in sore cases, Resembles Tirari Formation, Basal angular quartz granule layer, often

resting directly on underlying gypsum sediment,

Unit2, 025 LIMESTONE-GYPSUM. Greenish slightly sandy clay with 20 cm of interbedded thin

(0.5 coi) gypsum Jaminae at top, which grades laterally tnto laminated algal stem (tubules of

charophytes) limestone. The limestone and gypsum contain charophyte oogonia. The eypsum

contains scattered very coarse sand erains, and surfaces are assymetrically ripple-anarked, or

hare botryoidal “pulf” structure,

Unit |. 0,70 SAND. As for unlt 4. Orange and yellow stained, especially near base, greenish where un-

oxidized. Reworked distorted clay fragments from underlying units at base.

— Disconformity —

NAMBA FORMATION

3.28 SILTY CLAY, grey to black, tough. Grading down to grey, clayey, poorly sorted fine sand. Greasy

lustre on Jreguiar fracture surfaces. Gypsum patches and cracks at top infilled with overlying sand.

SUPPLEMENTARY SECTION, MILLYERA FORMATION

SECTION 6

EURINILLA FORMATION

At least 2.0 bright ced brown SANDS.

MILLYERA FORMATION

Unit?, 06.20 CALCAREOUS SANDSTONE. Very fine to medium grained moderately sorted sand,

30% carbonate, Grains pitted or frosted, subrounded to rounded. alternates with very fine

sand. Coarser sand contains charophyte tubules and rare oogonia, Some pink and black sand

erins, Fare carbonate grains, Weathered colour white (N10), unweathered greyish yellow

(5 ,

Elsewhere passes to hard platy limestone identical with 2. Impregnated with numerous white

fypsum cylindroids. Gradational contact with 6.

Unitf. 0.62 SAND. As for sand in 3 but uncemented, distinct contact with 5. Colour moderate red-

dish yellow (2¥7/4).

Unit5. 1.20 CLAYEY SAND. Maderntely sorted, with black and orange erains scattered through-

out, Jrregularly cemented into very hard massive nodules and sheets by fine grained white to

pink carbonate. Yellow and brown mottling common near base, white gypsum and carbonate

spots throughout. Yellowish grey (SY6/Z)-

Unit4+, 025 SAND, SILTY CLAY. Grades from clay to very fine sand, grains poorly rounded,

Colour yellawish (SY7/2) oxidized to moderate brown (SYR6/7) in patches.

Unit3, 0.5 LIMESTONE, CALCAREOUS CLAY. Varies Jaterally from burrowed soft calcareous

clay, with 30% silt to fine sand, to hard sandy white limestone, ‘The former bas 1-2 mm

diameter vertical burrows (insects?) and the latter has scattered charophyte oogonia and

shrinkage cracks. The base of the burrowed horizon is gradational, and lumps of the underly-

: ing unit are worked into it.

Unit2. 0.10-0.20 LIMESTONE. Laminated, platy, hard, metallic ring when struck. Constitutes

mimerous tubules of charophytes, and patches of “Coxiella", Contact with 4 not observed,

contact with 1 distinct, undulating.

Unit |. (0.50, On east side of channel, SAND. very fine grained, nodular white carbonete at lower con:

tact, Pale grey, Massive carbonate-cemented al top with shrinkage phenomena apparently fe-

tuted to drying. of carbonate. ;

On west side of channel. CLAYEY SAND. Moderate to well sorted, angular grains, Yellow-

ish grey (SY6/1) but speckled yellowish green, Grades up inte unit 3-

NAMBA FORMATION

0,10 Black tough clay, Sharp fiat upper contact.

EURINILLA FORMATION AND COONARBINE FORMATION TYPE SECTIONS; MILLYERA

FORMATION SUPPLEMENTARY SECTION

SECTION 7

Modern dune sands

— Disconformity —

COONARBINE FORMATION

Unit4. 6.70 SAND, very fine 1o medium grained, silly, Fine size dominant, well sorted. well

rounded, frosted. Light brown (SYR5/6), Al top is 20 cm of soft white carbonate, consisting

of 0.3 cm cylindroids and tubules (plant roots?) with }-2 cm lumps at the top, grading to

blotchy white carbonate as for unit 2.

186 R. A. CALLEN & R. H. TEDFORD

Unit3. 1,62 SAND, bimodal, medium-coarse atid very fine to fine. Bimedalily disappears downward,

grain size becomes finer, and sorting poorer, Some patches of white sand are present in the

essenually moderate vellowish orange (9YR3/6) coloured sequence. Top a4 moderate reddish

brawn (3YRS/6). Large scale cross-bedding is just visible. A well developed fossil carbonate-

rich 40) horizon marks the top, It is 50 cm thick and consists of moderately hard rather

irregular nodules and cylindroids, and gypsum cylindroids,

Uniz2 1.80. SAND, fine-grained ranging to coarse, grades down to CLAY-SILT, Moderately poor

sorting. No signs of stratification. Colour light brown to reddish yellow (6¥R5/6-3YRS/6)-

Weakly developed whitish carbonate patches ut top (soil horizon).

Unitl, 1.00 SAND. Bimodal: on medium-coarse grained aad very-fine grained boundaries. Dark red

brown (2YR4.5/6). Indistinctly horizontally laminated, upper contact sharp and flat, lower

contact obscure, apparently gradational,

— Disconformiry —

EURINILLA FORMATION .

Unit3 3,50 SAND, as above, Solour light brown (7YR6/4), Constitutes a single cross-bed set, Con-

tact with unit §, sharp, inclined, flat. Irregular gypsum as for unit £ at base, Upper 10-20 im-

pregnated with carbonate (1OYR7/3) of a fossil soil horizon.

Unie? 4.00 SAND, friable, fine to medium grained, bimodal. Course fraction well rounded,

dominant, colour light brown (6YR5.5/6). Constitutes a single cross-bed set, with low angle

cross-bedding. Contact with underlying unit sharp, undulating, cuts well down into unit 1,

Patches of very irregular tubules, nodules and cylindroids of gypsum occur at the top,

Unir}, O88 SAND, Medium @rained, sab-rounded to well-rounded grains with very fine grained

angular proportion (bimodal). Numerous coloured grains, opuques and biotite present. Silty

brown clay with gypsum forms pebble sized clasts, and clasts of underlying hmy sandstone

are present. Cross bed sets planar, 8-10 cm. Charophyte oogonia very common, and fragmen-

tal vertebrate bones present. White colour, Lower contoct erosional,

— Disconlormiry —

MILLYERA FORMATION .

1.1 LIMY SANDSTONE. Very fine to medium graincéd moderately sorted <lear-grained quartz sand

with 30-40% finely crystalline soft carbonale cement, Sand grains pitted of shiny, angular to sub-

tounded. Some grains of feldspar and ferruginous sandstone, flakes of hematite. Thin section shows

carbonate has recrystallized into radiating spherules, resembling some groundwater carbonates,

2.8 SAND, SILT, Silt to fine sand, 1-5% clay-carbonate matrix, forms strong cement by reason ot poor

eorting, of fremsgore Stains, Very poorly sorted with sub-rounded to very ungulur grains. Hard and

“white mottled,

SUPPLEMENTARY SECTION, EURINILLA FORMATION

SECTION &

Locality; CURNAMONA, Evrinille map sheets. Air photo ref: S. Aust. DepL Lands Svy, 161, run 3;

photo no. 4398. The section is situated on the west side of Tuke Pinpa, approximately 350 m north of the

only track crossing the Jake-

. . i Modern sand dunes

0,0-4,.0 SAND, Fine to medium grained, moderately sorted. SYR5/8, Strongly erosional base.

— Disconformity —

ICOGNARBINE FORMATION

0.30 SAND. Very fine to medium (averaging fine grained), poorly sorted, with sub-angular ta sub-

rounded polished or frosted grains (4¥RS/8). Erosional basal contact,

This unit may represent the Coanarbine Formation. It forms the basis of the longitudinal dunes.

— Disconformity — _

0.25 SAND. Very fine to fine, rather poorly sorted, clayey, Grains irregular, subangukar’, roleh, Colour

SYR4/7, Soft patchy carbonate well developed, With pipe like structure 5 cm diameter, This is a soil

horizon, and has a similar development &) those of the Furinilla Formation, The lithology is also simi-

lor hut there is a distinct contact at the base which appeared slightly crosional. The unit may be purt

ef ihe EURINILLA FORMATION.

EURINILI.A FORMATION

Unit3, 100 SAND. Clayey, very fine to fine, poorly sorted. Angular to sub-rounded frosted and

coyesely pitted sand grains, Colour 3YR5/8. Well developed secondary carbonate profites

constituting soft sandy pinkish white lime in lumps ard cylindroids 1-S cm across which

weather Gul. In lower part of sequence fractured himps 10-20 cm are common. The car-

honate profiles form numerous layers, concentrated toward the top of the unit, and represent

soil development (hence intermittent deposition is indicated}. }

Unit2, 5.33 SAND, Poorly sorted silty, slightly clayey fine grained, rounded (2.5YR4/8), Grades im-

perceptibly into overlying writ.

UNNAMED CONGLOMTRATE (?MILLYPRA FORMATION equivalent)

Unitl. O.8 Jnterhedde¢g CONGLOMERATE, SAND and CLAY. Consists of basal sand loosely

cemented with calcitim carbonate, cross-bedded on medium scule. Micaceous, At the base of

this sand bed ore granule size quan? ¢{prey, clear, dark grey, yellow), pebble size clasis of

ecey clay and subrounded calcareous orange-brown clay. Also large grains of brown perthite

feldspar! Maximtum grain size is 15 x 2 cm,

— Disconformity? —

CAINOZOIC ROCK UNITS 167

Overlying this is an uncemented medium to coarse sand layer, which is capped by coarsely

crystalline gypsum nodules and plates, weathering to a powdery crust.

The gypsum is followed by coarse grained to medium grained moderately sorted sand as

helow, with scattered very coarse grains. Grains are dull and subrounded to rounded.

The uppermost calcareous sand (5YR&/4) alternates with thin SILTY CLAY (10YR6/2)

with irregular patches of 5YR6/2 coarse grained very poorly sorted clayey sand. All con-

tact are distinct and flat. Apparently grades into overlying unit over short distance and by

intertonguing—no disconformity observed.

—Disconformity —

NAMBA FORMATION

Unit 8. 0.40 SAND. Very fine grained. Yellowish white.

Unit7. 3.85 CLAY. Puggy, soft, with powdery gypsum stringers at top. N5.

Unit 6. 0.85 CLAYEY SILT. Colour 5¥Y6/1 with 10Y6/2 patches.

UnitS. 2,23 CLAY. Hard, greasy, Colour SY6/1 (silty) to S¥4/1 becoming 5Y6/0.5 at base, Contact

with 4 gradational,

Unit4. 0.20 SILTY CLAY. Finely laminated, dark and bright orange brown siliceous limonite

nodules, dense masses of manganése oxide. Bright yellow green patches in clay. Basal contact

sharp, undulating, indicates an hiatus, Rare vertebrates.

Unit 3. 0.22 SAY, Dark olive (5¥4/1) greasy. Sharp basa] contact. Numerous vertebrates as for

unit 2,

Unit2. 0.12 DOLOMITIC CLAYSTONE. Pale green (SY8/2) with black patches. Subconchoidal

fracture, hard. Sharp contact with overlying units. Numerous vertebrates include lungfish,

crocodiles, turtles and marsupials.

Unitl. 0,18 CLAY. Light green.

In addition to this sequence, an erosional remnant of Coonarbine Formation is superposed on the top. of

the Namba Formation-Eurinilla Formation disconformity. The description is:

118 SAND. Yellow brown (7YR6/7) with prominent columnar jointing (15 cm rectangules) typical of

Coonarbine Formation. Fragments of Eurinifla Formation carbonate nodules occur at the base.

Also along the lake shore is reworked material from all the above units, forming outwash and aeolian

mounds of sub-Recent origin, This material is slightly older than the red brown dunes.

SUPPLEMENTARY SECTION, EURINILLA FORMATION

SECTION 9

Location: FROME, Coonarbine map sheets. Air photo ref.: S. Aust. Dept. Lands Svy. 395, Run 4,

Photo no. 9597. The section is situated on a cliff on the western edge of a small claypan on Eurinilla

Creek.

RECENT SAND DUNES

pees SAND. Very fine to fine grained, well sorted, with subrounded stains. Large scale dune cross-

bedding,

COONARBINE FORMATION

17-3.5 SAND. Clayey, up to medium grained, mostly fine grained, grains subrounded, Moderately

sorted. Weak thin horizontal bedding at base. Spotted with white carbonate patches, tending to 0.5 cm

diameter cylindroids near base {weakly developed soil profile), Colour 6¥YRS/7.

— Disconformity —

EURINILILA FORMATION

3.05 CLAYEY SILT. No sedimentary structure. Oxidized dark orange brown (SYR4/7)—original

colour (new patches) pale orange (10YR8/2). Gypsum beds several centimetres thick occur in lower

part, as for cap of 95 cm of massive disc shaped (0,5 cm) gypsum rosettes in red brown silt and white

gypsum flour.

NAMBA FORMATION ;

1.8 CLAYEY SILT. Very soft, 5¥5/0.5 with SYR4.5/4 patches.

— Disconfermity —

— Disconformity —

A NEW GENUS OF LATE PRECAMBRIAN POLYCHAETE WORMS FROM

SOUTH AUSTRALIA

BY M. F. GLAESSNER*

Summary

GLAESSNER, M. F. (1976).-A new genus of Late Precambrian polychaete worms from

South Australia. Trans. R. Soc. S. Aust. 100(3), 169-170, 31 August 1976.

New material indicates differences between Spriggina floundersi Glaessner and S.? ovata Glaessner

& Wade which are comparable with those between genera of living polychaete annelids.

Accordingly, a new genus Marywadea is proposed for ovata. The evolutionary significance of the

Sprigginidae is discussed briefly.

A NEW GENUS OF LATE PRECAMBRIAN POLYCHAETE WORMS FROM

SOUTH AUSTRALIA

by M. F. GLAESSNER*®

Summary

GLAgssNER, M, F_ (1976).—A new genus of Late Precambrian polychaete worms from South

Australia. Trans. R. Sac. S. Aust. 100(3), 169-170, 31 August 1976.

New material indicates differences between Spriggina floundersi Glaessner and S.? ovata

Gluessner & Wade which are comparable with those between genera of living polychaete

annelids. Accordingly, a new genus Marywadeu is proposed for ovata. The evolutionary signifi-

cance of the Sprigginidae is discussed briefly.

Introduction

The representatives of the Family

Sprigeinidae Glaessner (1958) are among the

most remarkable elements of the Ediacara

fauna from the Pound Quartzite of South Aus-

tralia. The Late Precambrian age, stratigraphic

posilion and geographic distribution of this

rock unit and its fauna need no further discus-

sion (Wade 1970; Glaessner 1971, 1972). The

arthropod-like appearance of Sprigeine is

attracting increasing attention (Cisne 1975, p.

61; Stanley 1976, p. 58). The reconstruction

of an ancestral crustacean by Hessler & New-

man (1975) shows startling resemblances with

Spriggina. Notwithstanding these, no canyinc-

ing evidence has been discovered which would

justify the transfer of the Sprigginidae from

Annelida to Arthropoda or prove a transitional

position of this family between two phyla. New

discoveries have, however, clarified and

emphasized the differences between the type

species §. foundersi and the species described

S.? ovata described by Glaessner & Wade

(1966). Its diagnosti¢ characters have at Icast

the same significance as those distinguishing

genera of living Polychaeta and for this reason

the following new genus Is proposed. It diflers

from Spriggina in all characters listed in the

diagnosis.

Taxonomy

Genus Marywadea nov.

fype species; Sprigginel avata Glacssner &

Wade 1966.

Fig. 1. Marvwadea ovata (Glaessner & Wade),

Latex mould of specimen from the Late

Precambrian Pound Quartzite of Ediacara,

§. Aust. x 2. (Outlines and surface slightly

distorted during fossilization; anterior

margin of the head pushed back causing

truncation of the outline and wrinkling of

the surface; some ventral structures may

be obscurely visible. Note that all other

specimens have smoothly curved anterior

outline and smooth surtace,)

* Centre for Precambrian

S, Aust. S006,

Research, Department of Geology, University of Adelaide. Adelaide.

170

Diagnosis: Prostomium half-moon-shaped, not

wider than the body with its appendages,

Integument thin, wrinkled and possibly show-

ing some underlying structures when com-

pressed, Body consisting of up to 5() short,

broad segments, occasionally with impressions

of bundles of long, curved setae. A pair of oval

impressions behind the prostomium suggests

the presence of teeth. The posterior end of the

body is broadly rounded.

Derivation of generic name: After Dr Mary

Wade who earlier expressed the view that

evata may be generically distinct from

floundersi; this has now been confirmed by

new finds.

Localities: Ediacara Hills, Brachina Gorge,

Bunyeroo Gorge, Mayo Gorge.

Number of specimens of M. ovata: 16.

Remarks

The Sprigginidae are not arthropods as the

head did not consist of the appropriate number

of appendage-bearing segments and the trunk

appendages are not distinctly jointed and end

in acicular setae. The mouth was. probably not

M. F. GLAESSNER

directed posteriorly, there was no labrum and

there is no evidence of antennae or a caudal

furca, On the other hand the head was con-

spicuous and relatively larger than in any

known annelid and its integument was more

strongly sclerotized in Spriggina (apparently

less so in Marywadea). There is evidence of a

simple pharynx in Spriggina and of two simple

teeth in Marywadea, suggesting relations to

Phyllodocemorpha; otherwise the Sprigginidae

are unhke living Annelida. Some evolutionary

advance in the direction of a primitive arthro-

pod is indicated, particularly in cephalization,

Ii may be parallel to the unknown evolutionary

lineage which had produced the two primitive

arthropods known from the Ediacara fauna

(Praecambridium and Parvancorina).

Acknowledgments

The specimen illustrated here was found by

Mr D. Westlake in August 1975 at Ediacara.

I am grateful to Mr J. Gehling, Murray Park

College of Advanced Education, who presented

to me casts and moulds of this and other

specimens.

References

Cisne, J. L. (1975) —Anatomy of Triarthrus and

the relationships of the trilobita. Fossils and

Strata (4), 45-63.

GLAESSNER, M. F. (1958) —New fossils from the

base of the Cambrian in South Australia.

Trans. R. Soe. 8, Aust. 81, 185-188.

Guarssner, M. F. (1971).—Geographie distribu-

tion and time range of the Ediacara Precam-

area fauna. Bull. venl. Soc. Amer. 82, 509-

GLARSSNER, M. F. (1972).—Precambrian palaéo-

zoviogy. Univ, Adelaide Cent. Precambrian

Res,, Spec. Pap, 1, 43-52.

Guarssnpe, M. F, & Wave, M. (1966).—The late

Precambrian fosils from Ediacara, South Aus-

tralia. Palaeontalogy 9, 594-628, pls 97-103.

Hessen, R. R. & Newnan, W. A. (1975) —A

trilobitomorph origin for the Crustacea. Fos-

sils aiid Strata (4), 437-459.

Staniey, 5. M. (1976).—Fossil data and the Pre-

eambrian-Cambrian evolutionary transition.

Am. J. Sci. 276, 56-76,

Wapr, M. (1970).—The stratigraphic distribution

of the Ediacara Fauna in Australia. Trans. R.

Soc. §. Aust. 94, 87-104,

VOL. 100, PART 4 30 NOVEMBER, 1976

TRANSACTIONS OF THE

ROYAL SOCIETY

OF SOUTH AUSTRALIA

INCORPORATED

CONTENTS

Robinson, A. C. and Smyth, M. E. B. The Vertebrate tee of a Vase

pelago, South Australia - - - - - 171

Shepherd, S. A. and Womersley, H. B. S. The Subtidal Algal and sesapank Ecology

of St Francis Island, South Australia —- - 177

Anstis, Marion Breeding Biology and Larval Development of Litoria herees

(Anura: Hylidae) - - - - - 193

Rowe, F. W. E. Restriction of the chiridotid genus Trochodota Ludwig (1891)

(Holothurioidea: Apodida), with the description of a new species

from South Australia - - - - - - - - 203

Annual Report of Council —- - - - - - - - - - 207

Award of the Sir Joseph Verco Medal _ - - - - - - - - 208

Balance Sheet - - - - - - - - - - - - 209

List of Fellows a i ey. (0

PUBLISHED AND SOLD AT THE SOCIETY’S ROOMS

STATE LIBRARY BUILDING

NORTH TERRACE, ADELAIDE, S.A. 5000

THE VERTEBRATE FAUNA OF NUYTS ARCHIPELAGO,

SOUTH AUSTRALIA

BY A. C. ROBINSON* AND M. E. B. SMYTH

Summary

ROBINSON, A. C. & SMYTH, M. E. B. (1976).-The Vertebrate Fauna of Nuyts Archipelago,

South Australia. Trans. R. Soc. S. Aust. 100(4), 171-176, 30 November, 1976.

The St Francis group of islands in Nuyts Archipelago was visited by a joint Royal Society of

South Australia and Fisheries Department of South Australia expedition in January, 1971.

Seven species of mammals, twenty-seven species of birds and sixteen species of reptiles are

recorded, together with comments on their habitat and abundance. The potential of the islands for

fauna conservation is briefly considered.

THE VERTEBRATE FAUNA OF NUYTS ARCHIPELAGO,

SOUTH AUSTRALIA

by A, C. Ropinson* and M, EB. B. Smytut

Summary

Ropinson, A. C, & Smyts, M. E. B. (1976).—The Vertebrate Fauna of Navis Archipelago,

South Australia, Trans. R, Soc. S, Aust. 100(4), 171-176, 30 November, 1976.

The St. Francis group of islands. in Nuyts Archipelago was visited by a joint Royal Society

of South Australia and Fisheries Department of South Australia expedition in January, 1971.

Seven species of mammals, twenty-seven species of birds and sixteen species of reptiles are

rycotded, together with comments on their habitat and abundance. The potential of the islands

for fauna conservation is briefly considered,

Introduction

The study of the fauna of islands can pro-

vide useful insights into. the biogeography of

the fauna on the adjacent mainland. Many

islands along the southern coast of Australia

were connected to the mainland whet: sea level

fell during the Pleistocene glaciations, and

samples of the coastal flora and fauna were

preserved on these islands as the sea level rose

during the interglacial periods (Main 1961).

With the accumulation of information on the

flora and fauna of these islands and the adja-

cent mainland, including the palaeofaunas and

floras, it may eventually be possible to recon-

struct the biological history of southern Aus-

tralia from the Pleistocene to the present. Fur-

ther, as information is assembled on the present

habitat preferences and tolerances of mainland

species, data on island faunas may assist the

construction of a palacochmati¢c history for

southern Australia. Another important aspect

of islands is their suitability for conservation,

Many of the islands around southern Australia

preserve relict populations of species that are

fare or extinct on the mainland. Studies such as

this should therefore contribute to the manage-

ment of these islands as fauna sanctuaries

muintaining these important populations,

This paper discusses the species of mammals,

birds and reptiles recorded from the St Francis

Island group of Nuyts Archipelago during the.

jeint Reyal Society of South Australia and

Fisheries Department of South Australia expe-

dition there in January, 1971, together with

some additional information gathered by sub-

sequent visitors to the islands.

The expedition visited four islands: St

Francis, Masillan, Fenelon and Dog. Mammal

trapping was carried our on St Francis [. (4

nights), Dog I. (1 night) and Masillon I. (1

night), Since then trapping hag been carried

out on Egg J. (D. Murray, pers. comm.), Data

were obtained from animals caught in traps or

observed by spotlight, from collection of bones

and from signs of mammal activity. Sherman

and Wire cage traps were used in trap lines for

a total of 153 trap nights (St Francis I. 105,

Dog I. 19, Masilion I. 19, Egg I. 10). Trap-

lines and spotlight surveys covered all vegeta-

tion associations on the islands and gave a

wide coverage of the areas. Specimens have

been lodged in the South Australian Museum

(SAM), registration numbers are cited.

The bird list is based on the observations of

all members of the expedition and compiled

by Mr P. Macrow. The records generally rep-

resent sight records, but where doubt existed as

to the identity of a species, a specimen was

shot for a positive identification,

The reptile list is primarily from collections

made on St Francis I. Only part of a day was

spent looking for reptiles on Dog, Masillon and

Fenelon Is.

* National Parks & Wildlife Service, Box 1782. GP.O., Adelaide, $. Aus. 5001,

+ Deceased; formerly of Department of Zoology, University of Adelaide, Adelaide, S. Aust, S000,

L72

MAMMALS

Mammals previously recorded [rom St

Francis 1. include the bandicoot Iscaden &be-

sulur naulicus und a ral kangaroo, presumed to

be a species of Betrongia, which had become

extinct by the 19203 (Wood-Iones 1924;

VYerco 1935), Three terrestrial and two marine

species are now added, and these are marked

by an asterisk in the Jist which follows. In addi-

tion, skulls collected on the island enabled

identification of the species af Betiongia.

Family PERAMELIDAE

Isoodon = obesulus (Shaw), SAM, M&8546-

M8549. Shori-nesed bandicoot. St Francis I.

A common animal preferring the grassy

areas on the higher parts of the island but also

occuring m the saltbush steppe association

covering the remainder of the island. These

bandicouts have survived the introduction of

cats and the conversion of a large part of the

island to grassland by cultivation. /. obesulus

remains common in south-eastern and south-

western Australia and there ure populations on

Kangaroo lL (Andrewartha & Barker 1969)

and Franklin 1. (Watts 1974).

Family MACROPODIDAER

Bettongia penicillata (Gray), SAM, M8353.

Brush-tailed betteng. St Francis 1.

Fragments of skulls were found in the suned-

hills behind Petrel Cove, but living animals

were not observed. Bettongs were reported to

be very common when St Francis Island was

first settled. WoodJories (1924) and Yerco

(1935) reported that the settlers introduced

cats to the isand to exterminate the bettongs

which were causing damage io vegetable gar-

dens. It seems likely thal alteration of habitat

may have also played @ part in their decline,

as this species nests in dense cover such as that

formerly provided by the sclerophyll shrub

community on the higher parts of the island.

The settlers completely destroyed this habitat

through clearance for wheat growing. 8. peni-

cillata was formerly widespread throughout the

southern half of Australia hut now appears to

he confined to southwest Westet'n Australia.

*Macropus cugenii (Desmarest). SAM, M&575.

Yaimmar wallaby, St Francis 1,

A single tooth row of this species was found

in the sandhills behind Petrel Cove, No living

animals were found, The settlers did not report

tammars on the island and it is possible that

they were either alreauly extinct, that the tndj-

A. C. ROBINSON & M. F_ B, SMY'LH

vidual collected wus introduced at some time,

or that it was left by sealers known to have

collected large numbers of wallabies on other

wlands (N. Wace pers. comm.). They were

formerly widespread on the south and south.

west Australian mainland and populations were

recorded from Kangaroo 1, Flinders 1. St

Peters [., and a number of Western Australian

islands, Today, in South Australia, tammuars

Temaiit common only an Kangaroo I. and on

Greenly 1. where they were introduced (Mit-

chell & Behrndt 1949); the St Peters 1. popula-

tion is extinct, the Flinders I, one is almost

eXtinet and the mainland population is reduced

to a remnant on Evre Peninsula (fF, Aitken

pers. comm-),

Family MURIDAE

*Rattus fuscipes (Waterhouse). SAM, M8541-

MBS45, M8598-M8600. Bush rat. Masillon and

Dog fs.

This species appeared toa be common on

these islands, where it nested in limestone

érevices and, possibly, mutton bird burrows. k

seems unlikely that the smaller islands of Nuyts

Archipelago cver supported mammals larger

than A. fesxcipes. Its absence from Ege Island

suggests that this island may be too small to

support @ population of R. fuscipes. We did not

sev or collect this rat on St Francis |, where

there wre extensive areax of suitable habitat,

hut again they may have been exterminated by

cats.

*Rattos rattus (Linn.), SAM, M8551. Black raz.

St Francis [

Two lower jaws of this mntroduced species

were collected in the sandhills behind Petrel

Cove, Na living animals were caught. A. ratty

was undoubtedly introduced by the early

settlers to St Francis I. and has since hécome

extinct.

Family OTARIIDAE

*Neophoca cinerea (Peron & Lesucus). Austra-

lien seu lion, Fenelon I.

This species visits all the islands and there

is it breeding colony on the beach of Fenelon

Island. In January 1971 this colowy numbered

approximately 50 jndividuals, including a num-

ber of pups. D. Murray (pers. comm.) pro-

vided the following estiraates of the size of this

colony in February 1973: mature bulls 7, pups

8, cows and immature bulls 36, The number

of individuals in the vicinily of the beach was

56-58

VERTEBRATES OF NUYTS ARCHIPELAGO

A South Australian National Parks and

Wildlife Service expedition in June 1975 was

table to land ot Fenelon T., but a count from

the boat showed mature bulls 4, pups 5, cows

and immature bulls. 3. They alsa noted a pos-

sible breeding colony of this species on Dog T.

Numbers recorded for this colony were bulls 3,

cows and immature bulls 10. In addition 20

sea lions were seen on Freeling, I.

*Arctocephalus forsteri (Lesson), New Zealand

fury seal. Fenelon 1.

The South Australian National Parks and

Wildlife Service expedition in June 1975 noted

40 fur seals on Fenelon 1. No evidence of

breeding was observed, They also noted 5 fur

seals on Freeling I.

BIRDS

No systematic list of the birds of the St

Francts Group has been compiled. The follow-

ing list contains comments on habitat and

abundance of birds observed during the 1971

expedition.

Family SPHENISCIDAE

Eudyptola minor (Stephens), Little penguin, St

Francis 1.

Common around the shores of Petrel Bay:

most of fhe birds were in a heavy moult.

Family PROCELLARIIDAR

Macronectes giganteus (Gmelin), Giant petret.

Dog I.

Beach washed specimen.

Puffinus fenuirostris (Temminck). Short-railed

shearwater. All islands visited except Fenelon.

Nesting burrows were found wherever suffi-

cient soil depth allowed excavation, Approxi-

mately one-third of St Francis I. was covered

by the burrows. Dunng the day most burrows

contained one adult bird and one egg in an

advanced stage of incubation. At approximately

20.00 hours each evening vast numbers of birds

returned to the island from feeding al sea.

Family OCEANTIDAE

Pelagodroma marina (Latham). White-faced

storm petrel, Dog and Fenelon Is.

Dried remains and wings were found, Small

burrows. on Fenelon JI. may belong to. this

species.

Family PHALACROCORACIDAE

Phakacrovorux varius (Gmelin). Pied

tnorant. St Francis I,

cor-

173

A small number of birds were fishing in

Petrel Bay and roosting in company with

black-faced cormorants at the eastern end of

thie bay,

Phalacrocorax fuscescens (Vieillot). Black-facea

cormorant, St Francis 1.

Approximately 20 birds roasted on the

eastern headland of Petrel Bay.

Family ANATIDAE

Cercopsis novachollandiae (Latham).

Barren goose. St Francis 1

Approximately 50 geese were observed and

flocks of 3 to 20 were seen feeding on the

eastern end of the island near the lighthouse.

The geese congregated around three small fresh

water soaks above granite boulders on the

eastern end of the island, Goose droppings

were ulsa found on Masillon I.

Anas sp. Unidentified teal. Egg I.

One bird was seen at sea near this island,

Family ACCIPITRIDAE

Haliaetus leucogaster (Gmelin). White-breasted

sed eagle. St Francis and Masillon Is.

Several adults and one immature bird were

observed fiying over St Francis I. and three

adult birds were seen fying over Masillon IL

Family PANDIONIDAE

Pandion haliaetus (Linn.). Osprey. St Francis 1.

One or two birds were observed on most

days at the eastern end of the island near the

lighthouse, Old nests of sea eagles or ospreys

were found on the eastern side of Dog I. and

the southern side of St Francis I.

Family FALCONIDAE

Falco peregrinus (Tumstall). Peregrine falcon.

Masillon I.

Only a single bird was observed,

Falco cenchroides (Vigors & Horsfield). Nan-

keen kestrel, St Francis, Dog and Masillon Is,

Several pairs on St Francis L

Family PHASIANIDAE

Coturnix pectoralis (Gould). Stvbdle quail, St

Francis I.

Several birds were flushed in the grassy area

near the lighthouse-

Family RALLIDAE

Rallus philippensis (Linn.}. Bonded landrati, St

Francis and Dog Is.

This species appeared common om St Francis

I. and two specimens were cailected. Only two

birds were sighted on Dog [,

Cape

174

Family HAEMATOPODIDAR

Haematopus fuliginosus (Gould). Seory oyster-

catcher. All islands visited.

A common bird of the rocky shorelines.

Family CHARADRIDAE

Vanellus miles wovaehullandiae (Stephens),

Spur-winged plover. St Francis 1.

Eight to ten birds were observed feeding

around the shores of Petrel Bay,

Charadrius cubricollis (Gmelin). Hooded dot-

tere]. St Francis I-

From two to ten birds were seen on the

beach in Petre! Bay each day.

Charadrius alexandrinus (Linw,), Red-capped

dotterel. St Francis T.

One bird was observed on the beach in

Petrel Bay in company with four Red-tecked

stints,

Family SCOLOPACIDAE

Calidris ruficottis (Pallas). Red-necked stint. St

Francis I,

Four to eight birds on the beach in Petrel

Bay, Onc specimen collected.

Family LARTIDAE

Larus noyuehollandiae (Stephens). Silver guil.

St. Francis f.

Nine to ten birds on the beach in Petrel Bay,

Larus Pacificus {Latham). Pucifie guif. All

islands.

A common bird of these islands. Adults and

immature birds wete present in about even

numbers. Approximately 20 birds furaged

along the shores of Petrel Bay,

Hydroprogne tschegrava (Lepechin), Caspine

tern, St Francis and Dog Is.

Two birds noted on each island.

Family PSITTACIDAE

Neophema petrophila (Gould). Rock parrot,

Found on all islands visited,

Numerous small flocks were flushed while

walking on St Francis J.

Famity HIRUNDINIDAE

Hirundo tahitica neoxena (Gould). Welcome

‘wallow. Found on all islands visited.

A common bird. Old nests. in houses, light-

house shed and caves,

Family MOTACILLIDAE

Anthus noyaeseelandiae (Ginelin). Pipit, All

islands visited,

Abundant.

A. C. ROBINSON & M. E. B. SMYTH

Faniily MELIPHAGIDAE

Meliphaga virescens (Vicillot). Singing honev-

eater. All islands visited.

A very common bird. Eight to ten birds were

present in the camp area at all times.

Family CORVIDAE

Coryus coronoides (Vigors & Horsfield), Aus-

tralian raven, St Francis L.

Common; » flock of approximately 30 birds

was observed as the expedition landed. Small

flocks were seen daily, foraging amongst the

mutton bird burrows. The lighthouse tower was

a favoured roost, and nest remains were found

here, Other nests were found on low. bushes.

REPTILES

Eleven species of reptiles are listed or men-

tioned for Nuyts Archipelago by Proctor

(1923), Waite (1923) and Worrell (1963),

Our expedition added another five species;

these are Indicated by an asterisk in the list

below. The islands of the group from which

each species is now known ig alse recorded.

Some contrasts between the abundance af

several species on St Francis and other off:

shore islands in the Bight are noted in the list

below. Possibly the drier climate of the Nuyts

Group is responsible,

Fumily GEKKONIDAE

“Underwoodisaurus mili (Bary], SAM,

R12858, R12863, R12870, RL2876, R12889.

St Francis, Masillon, Fenelon, Dog Is.

Common under limestone boulders er in

burrows in the sand by day.

‘Heteronotia binvei (Gray). SAM, 12878. St

Francis I,

Common under stones by day.

*Phyllodactylus = marmoraiux (Gray), SAM,

R12865, R12877. St Francis, Fenclon 1s.

Surprisingly uncommon, tor this is a very

abundant species on some other off-shore

islands. Found only under avolianite slabs on

exposed coastal arcas.

Family PYGOPODIDAE

Lialis burtonis (Gray), SAM, R12896. St

Francis L.

Only one seen during the visit,

Aprasia striolata (Lutken). Recorded fur St

Francis 1, by Kluge (1974); this is the speeci-

men referred by Proctor (1923) to Delia

fraseri, No Aprasia was collected in 197L.

VERTEBRATES OF NUYTS ARCHIPELAGO

Family AGAMIDAE

Amphibofurus fionni (Proctor). SAM, R12874,

Sc. Francis J,

Found only among the exposed granite

uraund the cdges of the island. Wooc-Jones did

not find it {Proctor 1923), but Worrell (1963)

has recorded it from St Francis f.

Family SCINCINAE

Hemiergis peronii (Fitzinger). SAM, R12862.

St Francis, Dog and Fenelon Is.

Surprisingly infvequently seen; like P. mear-

moratus, this is a common species on other off-

shore islands.

Egernia multiscutata (Mitchell & Behrndt).

SAM, R12857, RI2861, R12873, R128RS. Se

Francis, Dog, Masillon and Fenelon Is,

A very common species on the sandier parts,

burrowing under rocks or bushes and using

the mutton-bird burrows for quick retreats,

Lerista frosti (Zietz). SAM, R12859. Masillon

Another species usually common on off-

shore islands byt very scarce in the Nuyts

group.

*Lerista sp. (near picturata). SAM, R12880. St

Francis I.

A large member of the genus, with forelimbs

reduced to dimples and two toes on each hind

limb. Commonly found buried in sand under

stones.

“Menetia greyii (Gray). SAM, R12875, St

Francis I.

Rarely scen, probably because it js small,

quick and secretive,

Morethia obscura (Storr). St Francis t.

Again, rarely seen and yery difficult to catch,

ho specimens collected.

Tiliqua branchiale (Gunther). SAM, R12864,

RJ2879, St Francis, Fenclon Is,

Common in litter and around the buildings.

Family BOIDAE

Morelia spilotes. variegata (Gray). Carpet snake.

St Fraticis I.

178

Commonly seen in the morning and lata

afternoon, no specimens collected

Family ELAPIDAE

Drysdalia coronoides (Gunther), SAM, R12860,

Ri2881. White-lipped snake, St Francis, Masil-

lon and Fenelon Is,

Not frequently seen, but probably quite com-

mon,

Notechis ater (Krefft), SAM, RI2895. Tiger

snake. St Francis I.

Usually common on miutton-bird. istands, yet

only a few were seen even at night.

Discussion

The vertebrate fauna of the St Francis [.

group of Nuyls Archipelago is quite diverse

and remains Telatively undisturbed and rela-

tively free from introductions. The fauna of

most South Australian islands and indeed of

the mainland adjacent, is still incompletely

known,. and so biogeographical interpretations

are difficult at this stage. However, some com-

ments may be made on the importance of thesc

islands im conservation.

Island faunas are extremely vulnerable to

man’s interference, and the fate of Bettongia

penicillate. on St Francis J. illustrates this point

Tf further work there definiicly establishes that

this species rs extinct, the opportunity exists to

re-establish dense vegetation On the island and

introduce B. penicillate from Western Aus-

tralia. This should succeed as cats are no Jonger

present on the island. It is obviously a long-

lerm project but it merits consideration because

the island population could ultimately serve as

a reservoir of animals for release in suitable

areas of their former mainland range, In addi-

tion, it Rattus fuscipes is absent from St

Francis 1. it could be re-introduced from

neighbouring islands,

The majority of the birds observed are com-

mon on all islands in the area and on the

adjacent coast, but a notable exception ts the

Cape Barren goose, The total world population

at this endemic Australian species, although it

is increasing, is still dangerously low, and so

every attempt should be made to consetve the

known populations from disturbance.

References

ANDREWARTHA, H.G, & Barxer,, S, €1969).—In-

troduction to the atudy of the ecology of the

kangarca island wallaby Protemnodon engenit

(Desmurest) within Plinders Chase, Kan-

guroo Island, South Australis. Trans R- Soe.

S. Avst. 93, 127-132.

Kuuce, A, G, (1974).—A taxonomic revision of

the lizard family Pygopodidae. Misc. Publ.

Mus, Zool, Univ, Mich, 147, 1-221,

Main, A. R. (1961),—The occurrenee of Macro-

podidie on islands and its climatic and

ecologics|l implications.. J. Prac, R, Soc. W,

Aust. 44, 84-89,

176

MITCHELL, F. J. & BEHRNDT, A. C, (1949).—

Fauna and flora of the Greenly Islands, Part

I, Introductory narrative and vertebrate

fauna. Rec. S. Aust. Mus. 9, 167-179.

Proctor, J. B. (1923).—The flora and fauna of

Nuyts Archipelago and the Investigator

Group. No. 5—The lizards. Trans. R. Soc. S.

Aust. 47, 79-81,

Verco, J. C. (1935).—Jn B. C. Cotton (Ed.),

“Combing the Southern Seas”, chapter 8.

(Rigby: Adelaide).

Waite, E. R. (1923).—The flora and fauna of

Nuyts Archipelago and the Investigator

A. C. ROBINSON & M. E. B. SMYTH

Group. No. 10—The snakes of St Francis

Island. Together with a note on the name of

the geographical group. Trans. R. Soc. S.

Aust. 47, 127-128.

Watts, C. H. 8S. (1974).—The Nuyts Island bandi-

coot (Jsoodon obesulus nauticus). S. Aust.

Nat. 49, 20-24,

Woop-Jones, F. (1924).—“The mammals of

South Australia. Part II: The bandicoots and

the herbivorous marsupials.” (Govt Printer:

Adelaide).

Worreti_, E. (1963).—*‘Reptiles of Australia”.

(Angus & Robertson: Sydney).

THE SUBTIDAL ALGAL AND SEAGRASS ECOLOGY OF

ST FRANCIS ISLAND, SOUTH AUSTRALIA

BY S. A. SHEPHERD* AND H. B. S. WOMERSLEY}

Summary

SHEPHERD, S. A. & WOMERSLEY, H. B. S. (1976) .-The subtidal algal and seagrass ecology of

St. Francis Island, South Australia. Trans. R. Soc. S$. Aust. 100(4), 177-191, 30 November, 1976.

A subtidal survey of selected sites in the Isles of St Francis off the west coast of Eyre Peninsula,

South Australia, shows that upper and mid sublittoral zones similar to those of Pearson I. and West

I. occur. The upper sublittoral on rocky coasts is dominated by species of Corallina and Jania, with

Cystophora intermedia present near low tide level or sometimes as deep at 3 m. The mid sublittoral

is characterised by larger brown algae (Ecklonia radiata, Scytothalia dorycarpa, and species of

Cystophora and Sargassum), often with an understorey of red algae. The lower sublittoral zone

occurred between 47 and 57 m deep on the transect subject to greatest water movement, and is

characterised by red algae together with bryozoa, sponges and hydroids.

In the sheltered Petrel Bay- communities of the seagrasses Amphibolis antarctica and Posidonia

occur.

An algal species list is appended.

THE SUBTIDAL ALGAL AND SEAGRASS ECOLOGY OF

ST FRANCIS ISLAND, SOUTH AUSTRALIA

by S. A. SHEPHERD* and H. B. S. WoMERSLEY+

Summary

SeePxerD, S. A, & Womers.ey, H. B. 8. (1976).—The subtidal algal and seagrass ecology of

St. Francis Island, South Australia. Trans. R. Soc. S. Avst. 100(4), 177-191, 30 November,

1976.

A subtidal survey of selected sites in the Isles of St Francis off the west coast of Eyre

Peninsula, South Australia, shows lhat upper and mid sublittoral zones similar to those of

Pearson I, and West I. occur. The upper sublittoral on rocky coasts is dominated by species

af Corallina and Jania, with Cystophora. intermedia present near low. tide level or sometimes

as deep at 3 m. The mid sublittoral is characterised by larger brown ulgae (Ecklonia radiata,

Scytothalia dorycarpa, aud species of Cystophara and Sargassunt), often with an understorey

of red algae. The lower sublittoral zone occurred between 47 and 57 m deep on the transect

subject to greatest water movement, and is characterised by red algae together with bryozoa,

sponges and hydroids.

In the sheltered Petrel Bay, communities of the seagrasses Amphiholis antarctica and

Posidonia occur,

An algal species Jist is appended.

Introduction

‘The marine flora of the Great Australian

Bight is little known, Apart from the intertidal

tegion, which has been briefly discussed by

Womersley & Edmonds (1958), the subtidal

region is known only from various drift col-.

lections and the ecological account of Shepherd

& Womersley (1971) of Pearson Island,

towards the eastern limit of the Bight.

Ao expedition to the isles of St Francis,

lying off Ceduna, supported by the then

Fisheries and Fauna Conservation Department

and the Royal Sweiety of South Australia,

Visited the islands from 4-11 January, 1971.

This provided the opportunity for a brief sur-

vey of the subtidal ccology of selected sites

subject to varying degrees of water movement.

Although limited in time, these studies provide

the first such information from the northern

part of the Great Australian Bight.

The isles of St Francis comprise nine small

islands, the largest of which, St Francis. I.

(Fig. 1) is about 4 km across and lies at

32°9U'S, 133°18'E, about 56 km from the

mainland, The islands are granitic, rising

steeply from the sea-floor, and subtidally the

topography consists of massive blocks and

sheets of rock. In sheltered areas (e.g. Petrel

Bay), the sandy sea-floor slopes more gently

into deeper water.

The short stay on the island prevented a

detuiled survey, but three survey sites were

chosen on St Francis I., subject to different

degrees. of water movement. One site was

chosen on nearby Masillen L, and collections

were also made at Egg I.

The field work was limited to the subtidal

region, but brief observations of the intertidal

region indicated that the organisms and zona-

tion present were similar to those described by

Womersley & Edmonds (1958) for such grani-

lic steeply sloping coasts,

Methods

The following transecls (Fig. 1) were

chosen, and in cach case the transect ran nor-

mal to the coast, from low walter level down

the slope te the depth where rock was buried

by sand, except for transect D which was pre-

dominantly on sandy bottom.

* Department of Fisheries, Gawler Place, Adefuide, §. Aust. 5000.

t Department of Botany, University of Adelaide, Adelaide, S; Aust, 5000.

178

St Francis 4,

Masillon | 3P aa

Fig. §. Map of four of the isles of St Francis

showing the position of the four transects

(A-D). Inset shows the situation of the

islands in the northern Great Australian

Bighi.

Transect A was on the NW corner of St

Francis I., under conditions of strong water

movement, and terminated at 57 m depth. A

sampling gap (32-38 m deep) in this transect

was filled by collections from a similar site on

Ege I.

Transect B was on Masillon 1, subject to

moderate water movement, and terminated at

33 m depth,

Transect C on St Francis 1, was aubject to

relatively slight water movement, and des-

cended to 20 m.,

Transect D in Petrel Bay on St Francis 1.,

wus the most sheltered site, with a gently slop-

ing, sandy, sea-floor dominated by seagrasses-

S. A. SHEPHERD & H. B. 8. WOMERSLEY

On transects A, B and C, the diver first

swam aboul 2 m above the bottom along the

transect, estimating the percentage cover and

recording the vertical range of the prominent

brown algae. This was then repeated on twa

parallel lines, one each side of the first transect

and about. 15-20 m distant. Estimates of cover

for particular depths are given as the average

of these three valucs, which were made sub-

jectively as a percentage on @ scale of 0-10,

Many species vary considerably in percentage

cover over a horizontal distance of some

metres, but the figures given provide an overall

assessment of the cover along the transects.

Communities were recognised by the upper

stratum dominants, this being the most satisfac-

tory method on such surveys. At the depths

studied. alyac were dominant except at 30-57

m on transect A, and within each community

quantitative samples of the upper stratum were

taken for biomass estimates. This was done by

counting the plants in a hoop af § m* area,

placed sequentially along a horizontal line

some 16-24 times, so as to give a total

sampling area between 2 and 3 m®*. and the

humber of plants per m? calculated. The aver-

age weight of an individual plant was deter-

mined by weighing a random sample of ID

plants, and the biomass per m= then calculated.

The other strata were sampled by means of 4

to 8 sequential samples, each of $ m*,

On transect D in Petrel Bay, a diver was

towed on an underwater sled behind a boat,

and the distribution and depth range of the sea-

grusses Were noted, and photographs taken.

‘The algal samples were preserved in 4%

formaldehyde-sea water, and taken to the

laboratory for determination and analysis. Bio-

mass figures are based on the wet weight of

the preserved collections after removal of sur-

face water, These estimates should be taken as

examples only of the size and variation in bio-

mass of the community dominants, since the

restricted diying time on a short expedition

such as this limits the range of transects, and

the number of samples that can be taken, The

transect samples were supplemented by other

géneral observations and collections, and by

photography.

Depths were measured by capillary and

mechanical depth gauges, and the results aver-

aged and adjusted to approximate low lide

level.

Environmental factors

The shart duration of the survey precluded

Getailed studies on environmental factors, but

SUBTIDAL ALGAL, AND SEAGRASS ECOLOGY ivy

the following information is available. The

isles of St Francis rise from a maximum water

depth of about 60 m at the southwest of St

Francis I. and are comparable to the Pearson

Islands in their distance from the mainland and

in their topography,

Water movement

St Francis I. is subject to a strong south-

westerly swell of 10-12 second period, prevail-

ing throughout the year, similar to that at

Pearson I. and West I. (Shepherd & Womers-

ley 1970, 19714). In summer, a short southerly

swell is generated by the strong southerly winds

which blow for about 12 hours each day and

are characteristic of this part of the Great Aus-

tralian Bight. Wave action on all parts of the

islands facing south to west is strong in the

intertidal region.

Temperature, salinity and nutrients

Sea surface temperatures range from

18-20°C in summer ta 14-15°C in winter,

according to. Vaux (1970) and data obtained

from various oceanographic stations in the

Vicinity (C'S.I.R.O, 1967a, 1967b, 1968, 1969).

In summer, bottom temperatures at 50 m depth

are 2-3°C lower than sea-surface temperatures,

During the study, the surface temperature was

18°C off the island and about 20°C inshore in

Petrel Bay.

aot

Q SPONGES

C HYDROLDS

Salinity, phosphate, nitrate and oxygen levels

are similar to those for Pearson L., viz. salinity

35.6-36.2%-«: inorganic phosphate 0,09-0.17 pg

atom/ litre; nitrate about 0.3 pg atom/litre; and

oxygen saturation 93-103 %.

Submarine light intensity

Light penetration was not measured, but

according to H. Jitts (pers. comm,) it corres-

ponds to that for Type Il oceanic water of

Jerlov (1968), and is thus only slightly less

clear than the waters about Pearson T.

The algal and seagrass ecology

The algal-dominated subtidal photic zone at

other localities in South Australia has been

found to present three main zones, designated

as the upper, mid, and lower sublittoral zones

(Shepherd & Womersiey 1970, 1971). These

zones are alsa apparent in the areas studied at

St Francis I.

Communities of the rocky coast will be des-

cribed first, including these on both horizontal

and sloping rock but not those in crevices or

under overhangs, followed by the sea-grass and

algal communities of sheltered, sandy areas.

The communities studied are essentially those

subject to suflicient light intensity to be plant

dominuted, but prominent animal species ate

mentioned where present.

m BALANUS NUGRESCENS

a “Fo INTERMEDIA

$F vvarovesen HARVEVANUM

SCYTOTHALIA DORYCARPA

EARGASSUM BRACTEOLOSUM

ECKLONIA RADIATA

sa RED ALGAE

% ZONARTA

P BRYOZOA

Fig. 2, A vegetation profile of transect A (St Francis L)-

mid S.L.

S$. A. SHEPHERD & H. B. 8. WOMERSLEY

ora CORALLINA

CYSTOPHORA PELL INATA

f CYSTOPHORA MONILITORHIE

Fig. 3. A vegetation profile on transect B on Masillon I. Sce Icgend on Fig. 2 for other algal taxa.

a UUJANIA

mid S.L.

*¥ CYSTOPHORA MONILIFERA

a OSMUNDARTA PROLIFERA

SARGASSUM

VERRUCULOSUM

SARGASSUM

¥ BOTRYOCLADIA GBOVATA

Tenereawens:

Fig. 4, A vegetation profile of transect C on St Francis I, See legend on Fig. 2 for other algal taxa.

A. ROCKY COASTS

Transects A, B, and C traverse rocky areas,

generally siceply sloping and imcluding both

horizontal and sloping rock surfaces. The

marked light gradient with depth, coupled with

the. considerable gradient in water moyement

within each transect (especially A) and also

between the transects, gives rise to a fairly dis-

tinct zonation of algae. Profiles for transects

A, B, and C are given in Figs 2-4, and the

depth relationships of the communities to water

movement are given in Fig, 5.

L. Upper sublitteral zone

This zone is subject to the most intense water

movement, and varies in vertical width from

5(-7) m on rough-water coasts (transect A,

Fig. 2) to 2(-3) m on sheltered coasts (transect

C, Fig, 4). Communities of this zone typically

have a single, dense, stratum of fairly uniform

height, ranging from 15-20 cm for the Cysto-

phora intermedia community to 2-3 em for the

Jania community.

Corallina cuvieri, in high-light situations (i.e.

especially horizontal surfaces and those facing

SUBTIDAL ALGAL AND SEAGRASS ECOLOGY ik]

TABLE f

Species aud biomass (g/m) compostrion of upper sublittcral

communities tn samples taken at about 1(-1,5) m depth on

transects A, B and C. “PY indicates xpurve occurrence

althwugh net present in sample.

‘Transect A B c

Ara sampled (m=) O25 1.25 637

Water movement Strong Moderate Slight

Dominant Species

Cusophora intermedia (,280 p 30

Myrivdesma harveranum 1,400

Corallina cuvierl

1, eringaia 2,800 4,200

Janie favrintatsa Pe

Other Species

Canterpo brownli 70

Caulorpa papillon 140

Cyrophera gracilix <i

Lobosnita blenspldata 40 ay

Pachydielyon panioilatam Str Xr su

Saryax\urm spp. <10

Callophyllis raralfreiray 28) 15

Champiu obsolpta 40

Dasva clavigera <0

Griffithsia tages 260

Hypnea sp, 40

Laurencia filifarmis £.

herernclada 290

Séapara harveyiana <i0

Polvaiphonia nigrita <i

Total coverage 100 100 lub

Biomass. g/m! 3,950 2,840 4,330 [1,900]

Number of species 14 4 5

In each case the biomass value of the species charactens-

ing the community is in bold type,

* This sample was taken from a distinct Juala community

ala depth of about 0.5 m (see Fim 4).

nerth or cast) and subject to strang to moder-

ate water movement, forms an almost pure

community completely covering the rock sur-

face, In calmer areas a Jania fastigiafa com-

munity, presenting, a somewhat similar aspect

of short, tufted plants, replaces the Corallina.

The Cerallina cuvieri community extends up-

wards into the lower eulittoral zone of the

intertidal in rough-water situations, as des-

cribed by Womersley & Edmonds (1958, p.

232),

Cystophora ihtertiedia forms a fairly pure

cammunity under slightly less extreme water

movement than Corallina, and also on sloping

{rather than horizontal) surfaces subject to

somewhat lower light intensity. While Cysto-

phora intermedia may be dominant in. such

situations, in lower light intensity Myriedesmie

harveyanum becomes co-dominant, with

numerous associated species of green, brown

and red algae (see Table 1).

Cystophora intermedia is rare within the

Corallina cuvier? community, but may be com-

STRING WATLR MOVEMENT G&ADLENT SLICK!

TRANSECT A. 6 c

0 7

WRIGDESM

HARVEFANUM

‘

a. (VST. MONILTFORMLS }

S SANG. BRALTEUSUSIIF ¢

CYST POWILIELRA

SARG. VPRRUCELOSUM

cyst. PRCTUMATA

n> 1

Sy et

ESKLINLA RADIATA

SLPTOThALLA DORYEARCA

Fig. 5. Change in vegetation patterns along water

movement and depth gradients.

mon near the upper and lower boundaries of

this community. Its occurrence at the upper

limit (i.e. near low tide level} agrees with the

observations of Womersley & Edmonds (1953)

that it marks the sublittoral fringe. but at St

Francis 1. it is not confined to this zone, occur-

ting also as decp as 3. m,

2. Mid sublittoral zone

As at West I. and Pearson I[., this zone on

St Francis I. is characterised by larger brown

algae 30.cm=1 m in height, forming an upper

canopy or stratum over a lower stratum mainly

of red algac 5-25 cm in height. The upper

limit of this zone depends on the intensity of

water movement as described tor the upper

sublittora) zone, and the lower Jimit on the

limiting depth of large brown algae; this ts

about 45(-47) m deep on transect A. The

vevetation profiles of Figs 2-4 represent the

appearance of this zone on transeets A, B, and

C and the relations of the vegetation patterns

with waler movement are shown in Fig. 5. The

average cover of the important upper stratum

species is given in Fig, 6.

Several communities could possibly be recog-

nised in (his zone, but more extensive studies

than were possible in the time available ure

needed to establish their validity. The domin-

ants and understorey Species will therefore be

discussed more generally.

Ecklonia radiata and Scytothalia dorycarpa

(Fig. 9) dominate this zone under conditians

of considerable water movement at the rough-

182 S, A. SHEPHERD & H. B. S. WOMERSLEY

TABLE 2

Biomass (g/m*) composition of mid. sublitioral species in samples taken at certain depths on 3. transects. Further

data on the vertical range of the species is given in the appendix

Transect A B

Depth (m) 6 13 35 6 22 32 6

Area sampled (m2) 0.5 1 1 05 0.5 0.5 0.5

Upper stratum

*Ecklonia radiata 2,200

* Scytothalia dorycarpa

Cystophora pectinata

Cystophara moniliformis

Myriodesma harveyanum

Sargassum bracteolosum

Sargassum varians

Sargassum verruculosum

Sargassum linearifolium

Sargassuni héteromorphum

Sargassum decipiens

Cystophora brawnii

Cystophora subjarcinata

Cystophora monilifera

Upper stratum coverage (9%) 100. 95 35 90 50 60 80

Upper stratum biomass

(g/m) 4,650 3,775 2,210 2,250 2,460 3,700 3,250

Lower stratum

Brown. algae

Dictyopteris muelleri

Dictyata diemensis

Dictyota prolifera

Chlanidophora microphylla

Glossophora nigricans

Hydreclathrus elathraius

Lobospira hicuspidata

Pachydictyon paniculatum

Zonaria spiralis

Zonarta sinclairii

Zonaria inrneriana

Red algac

Austraphyllis aleicornis

Ballia callitricha

Botryocladia obavata

Chamipia affinis

Cliftonaea pectinata

Delisea hypneoides

Delisea pulchra

Kallymenia cribrosa

Laurenvia filiformis f,

dendritica

Laurencia spp.

Osmundaria prolifera

Plocamium angustum

Plocamium cartilagineum

Plocqminm mertensti

Placamium preissianum

Pterosiphania sp,

Rhodophyllis membranacea

Sonderaphycus australis

Webervanbossea kaliformis

1,800 450 900 -1,200

200 50 1,380 = =2,500

PTILItbaitias

Pitt dd

Privy ddd

wasuu | ro | Sus

TTT |

$18!

105 3

TTT ttt

lStr ltt

Pll bl td

LLEtbti dd

| |

25 180

680

— 50

ee:

— 50

1160 1,610 150 1,040

Ll Ll peal | |

lSllSslusil |

Ll td) aseles

L| |) Sell] |

i —

Blulaulwussws||S

Biomass

Pld td

all lwel Sl |

215

| | inns | |

Bes

ur | | laal aS& |

1,214

3,615

Total number of species

in Sample 4 7 25 24 22 12 11

Total Biomass. 4,650 3,845 3,098 3,410 4,070 3,850 4,290 6,020 4,151

* Values for Ecklonia and Scyrothalia are mean values over 2-3 m2,

4,445

SUBTIDAL ALGAL AND SEAGRASS ECOLOGY

UPPER STRATUM

LOWER STRATUM

meee enaee

WASOTININYYAA WASSYONWS

SNYIYVA WNSSVONVS

SNAId154a0 WOSSVDuVS

jo WNSQ10dLI vad WNSSVOXVS

(n> VAVNTDuVAENS YHOHAOLSAD

VLVNILIAd YYOHAOLSAD

SIWHOSTTINOW YHOHdOLSAD

VHSSITINOW VHOHdOLSAD

é ~>~ [va]

VIVICVY VINOTNDS

cover of prominent upper stratum species of the mid sublittoral zone, and total % cover of upper and lower strata

Fig. 6. Vertical distribution of %

on transects A, B and C.

183

IE4

water site (transect A, Fig, 2), but with Jess

water movement (trunsect B, Fig. 3) other

Species of brown algac (Sargassuni Aracren-

lexeat and C'ystuphora moniliformis) also be-

come prominent, With greater shelter (iransect

C, Figs 4, 7, 8), Cystophora inenilifera, C.

pectinata and Sergasswn vyerrucuiosunt are

gassum decipiens and S. varfans. These species

are most common in the upper part of the mid

sublittoral, with Eeklonia and Scylotheelie still

common in the lower part of thts zone (Fig. 4).

Sargassum Bracteolasum and Cystophord

menififornis on transect BL and §, verruen-

lovuen and C. nnartilifera on transect C, have

similar vertical distributions, cnabling Feld

recognition of algal subzunes dominated by

these species pairs.

Unuetstorey species are spatse over much

of transects A and C, and moderately common

only on transect B where the canopy ts less

dense. The distribution of many understorey

species is too patchy to show any obvious rela-

tionship with either the distribution of upper-

stotey species or with apparent environmental

factors.

The commonest understorey species are of

Plocamivm (P. angustam, Po mertensi and P.

preissionium). They occurred on all] three tran-

secis, with a vertical depth range of as much ay

50 m; where they are rare or absent on hori-

zontal surfaces, they are usually present on

vertical ones.

Other species with a wide vertical range are

Caulerpa Srownij, Lebospira bicuspidate and

Pachydictyon panicularuni; these species are

known to he tolerant to a wide range of light

intensity and of water movement. Some other

species [e.g, Gloxsophora nipricans, Austro-

phivilis aleicornis, Cliffanaea pectinate and

Delisea Aypneoides) were found only in Geeper

water; most of these species occur also at West

T. and Pearson L, with similar distribution an

conditions of lew light (and slight water move-

ment at depth}.

On transect C at 17-18 m depth (Fig. 4),

Where os an abrupl decline in the number of

upper stratum species and their coverage, and

aim increase in coverage of several species of

the lower stratum, e.g. Osinutdaria prolifera,

$3. A. SHEPHERD & H. B, S. WOMERSLEY

Borryocladia obovata and Hydroclothris clath-

rains, This community forms a band [2 m

wide lying immediately above the sandy bottom

at about 20 m depth, and the species are appa-

rently tolerant of the sedimentation which is

Pronounced over this narrow band,

The cover of upper and lower stratum spe-

cies, with depth, is given in Fig. 6, Upper

stratum cover is highest between 5 and 15 m

depth, declining with depth, whereas lower

stratum cover is lowest where the upper stra-

tum 1s most dense, and in general increases

with depth until light becomes limiting.

3. Lower subliteral zone

Only on uunsect A does rocky substrate

descend to sufficient depth for the lower sub-

littoral zone dominated’ by red algae (Shep-

herd & Wamersley 1970, 1971) to occur. On

this transect, a communily (Fig. 10) of red

aleac together with bryozoa, sponges and

hydroids, occurs between 47 and 57 m deep,

The community is rich in algal species (but of

low biomass), the most common being Plo-

cuniion ampgustim, P. mertensii and PF. preis-

Signum, several other species (Rhodymenta

australis, Gattva pinnella, Rhedocallis elegans

and Kallymenia spinosa) were found only in

this collection. Algal cover in this community

is low, averogmg 10% (5-15%), indicating

that 57 m is close to the depth (i.e. Light) limit

for most algae in this region.

B, THE SEAGRASS COMMUNITIES IN

PETREL BAY

Three seagrass communities occur in this

sheltered bay, forming bands around the bay

dependent on substrate ani depth.

Amphibolis antarcsica fringes the shore from

low walter mark to U5 m below, attached by its

rhizome-root system te calcarcous recfs of low

relief. Below these reefs the bottam is sandy,

and at a depth of about 2 m, Posidema osten-

feldii forms a fringe community about 20 nm

wide around the hay, Beyond this, descending

to 22 m decp, Posidonia australiy (narrow leat

form} i dominant in fairly continuous beds,

Beyond abqut 22 m deep, P. australis hecornes

sparse, and at the ume of the survey a loase-

lying but apparently healthy community ol the

red algo Hennédya crispa occurred at this

depth.

Fig. 7, Algal community at § m depth on transect C, Note Cystaphora mronilifera (top sett), Scylo-

thalia doryearpa (tep cight) and several species of Sarvaysum (ventre and lower righr)..

Fig. &. Algal community at 10 m depth at transect C. Note species of Sargassiem (centre left), Eckloptia

radiata (top right) and €ysropherd moniliferd (contre and lower eight),

185

SUBTIDAL ALGAL AND SEAGRASS ECOLOGY

S. A. SHEPHERD & H. B. S. WOMERSLEY

SUBTIDAL ALGAL AND SEAGRASS ECOLOGY 187

Discussion

Algal zones within the sublittoral, and the

distribution. cover and biomass of the com-

ponent spectes, have been described for many

Coasts elsewhere in the world. Recent accounts

are those of Liining (1970) from Heilgoland,

Boudouresque (1971) from the Mediterranean,

and Mann (1972) from the Atlantic coast of

Canada. These and other accounts show that

broad algal zones, correlated with light inten-

sity and the degree of Walter movement, occur

in the photic zone on most coasts,

Although limited in extent, this survey of

the subtidal algal vegetation of St Francis

I. shows a similar zonation pattern to that

at West [ and Pearson 1, (Shepherd &

Womersley 1970, 1978). As at these islands,

the vertical extent of the upper sublittoral zone,

and to a lesser extent the mid sublittoral, is

dependent on the degree of water roughness

(with whrch light penefratton is also associa-

ted), The extent of the upper sublittoral zone:

probably corresponds with the depth to which

“white water” (1.8. turbulent walter carrying air

bubbles) penetrates under average swell condi-

tions. Ried] & Forstner (1968) considered the

vertical height of their “inner surf zone”

{Riedl 1971) ta correspond to 2.5 x wave

height, and this could also be applied to the

upper sublittoral zone on South Australian

coasts Where wave heights are 1.5-2 m in a

moderate swell. Chapman (1967) in discussing

the presence of a suhblittoral fringe in many

parts of the Pacific is largely referring to this

upper sublittoral zone. The term “sublittoral

fringe” is best restricted to the zone emergent

during suck back of waves at low tide, when

this zone is ecologically distinctive ( Womersley

& Edmonds 1952)..

The mid sublittoral zone at St Francis I.

shows similar features to this zone at West DL

and Pearson I., being dominated by the larger

brown algae and with an (inderstorey of mainly

red algae. Further studies muy show chat dis-

unet conimunities could be recognised in this

zone, since competition between the various

dominant species is apparent, and, over the

considerable depth range, both light intensity

and degree of water movement vary consider-

ably. While most species show typical “bell-

shaped" distribution patterns (as discussed by

Whittaker 1967), some (e.2, Cystophora moni-

lifera, C. subfarcinata) apparently show

slightly bimodal distributions (Fig. 6), pro-

bably due to competition with other species

better suited to the environment within their

extremes.

The lower sublittoral zone of red algae was

observed only in depths of 47-57 m at St

Francis [., corresponding to the situation at

Pearson E. rather than wt West I, and reflecting

the ¢larity of the water. This zone lies below

the light intensity necessary for the larger

brown algac and grades to the flower photi¢t

levels of the red algae. At St Francis L, inter-

mixing of lower sublittoral red algae with

fauna such as hryozoa, sponges and hydroids,

was More prominent than at West 1,

Although sublittoral zones are well defined

at St Francis [., this characteristic is empha-

sized by choice of transects on steeply sloping

shores involVing steep light and water move-

ment gradients. On more irregular shores. dis-

linct zonation is less apparent.

Apart from ecological differences associated

with depth, which reflect mainly the decrease 1

light intensity, light relationships are apparent

in the mid sublittoral zune where a dense upper

canopy may reduce the light reaching the lower

stratum by up to 95%. This effect was well

shown on transect A at 5-15 m depth and

transect C at 7-13(-15) m depth, where a

dense canopy covered a sparse understorey.

Where a dense canopy exists with considerable

water movement, reduction of the understorey

may alsc be due to the physical effect of the

larger fronds sweeping over the rock.

The effect of sediment (fine sand or silt

stitred up in stormy weather and settling on

the seabed under calmer conditions) was evi-

dent in tLyo places. Near the end of transect B,

at about 30 m depth, sediment is present on

rocky surfaces and here there is an abrupt

decline jn cover of the lower ‘stratum, At the

end of transect C at 18(-20) m depth, where

sediment also covers the rocky bottom, there

is a distinctive community of certain red algae

(Rotryocladia ebovate, Osmundaria prolifera)

which can tolerate sediment, The effect of sede

Fig. 9. Algal community at 16 m depth on transect C. Note Ecklonia radiate {top right and lower left)

and Seyterhalia dorycarpa (centre)-

ig. (0. Sparse red algal community at 57 m depth on transect A,

188 5. A. SHEPHERD & H. B. 8. WOMERSLEY

ment in inhibiting algal colonisation and

growth has been recently discussed by Grigg &

Kiwala (1970)-

The survey of St Francis I. was limited in

time, the area covered, and in the variety of

habitats sampled. Nevertheless, the subtidal

algal flora appears fairly rich. with some 138

species recorded, compared to 160 for Pearson

1. and 132 for West I. Further studies would

certainly extend this number considerably,

Appendix: Algal species fist

Identifications are by H. B. S, Womersley, Dr G. T. Kraft (Mychodeaceae, Dicranemaceae and Acro-

tylaceae) and Dr BE. M, Wollaston (Crouanieac).

CHLOROPHYTA

Caulerpales

Caulerpa brownit (C.Ag.) Endlicher

Caulerpa cactoides (Turn.) C. Agardh

Caulerpa flexilis Lamouroux

Caulerpa flexilis Lanjouroux var. muellert

(Sond.) Womerslcy

Caulerpa hedleyi W. v. Bosse:

Caulerpe longifolia C.Ag_{. erispata (Harv.)

Womersley

Caulerpa obscura Sonder

Caulerpa papillosa J, Agardh

Cuulerpa scalpelliformis (R.Br.) C, Agardh

Caulerpu simpliciuscula (Turner) J. Agardh

PHAEOPHYTA

Dictyotales—Dictyoteae

Dictyota diemmensis Knetzing

Dictyota furcellata (C.Ayg.) J. Agardh

Dictyota prolifera Lamovurowx

Dilophus fastigiatus Sonder

Dilephus robustuy (1.Ag.) Womersley

Pachydictyon paniculainm J. Agardh

Pachydictyon nov. sp?

Glossophora nigricans (J.Ag.) Womersley

Lobospira bicuspidala Areschoug

Zonariewe

Chlanidephora microphylla (Harv.) J. Agardh

Dictyapteris muyelleri (Sand.) Reinbold

Lobophore variegata (Lamx.) Womersley

Zouaria crenata J, Agardh

Zonaria sinclairii Hooker & Harvey

Zonaria spiralis (J.Ag.) Papentuss

Zohkaria turneriang J, Agardh

Nov. gen?

Chordariales—Chordariaceae

Corynophlaea cystophorae ¥. Agardh

Bactrophora filum (Harv.) J. Agardh

Bactrophora vermicularis J. Agardh

Polycerea nigrescens (Harv. ex, Kuetz.) Kylin

Sporochnales—Sporochnaceae

Bellotia ériopherum Harvey

Sporochnus comosus C. Agardh

Dictyosiphonales—Giraudyaceac

Giraudya spiacelarioides Derbes & Solier

Punctariaceae

Hydroclathrus clathratus (C,Ag.) Howe

Laminariales—Alariaceae

Ecklortia radiata (C.Ag.) J. Agardh

Fucales—Cystoseiraceae

Seytothalia dorycarpa (Turn.) Greville

Cystophora brownii (Turn.) J. Agardh

Cystophora intermedia J. Agardh

2-38; B, 4-7; C, 2-6, 19

Ud we

‘S

, 13-18: C.19;D,2

BC, 10-13

P>PuUy Se >>>

Ka Pb b> lA

ieee

, 32-38; B, 636, 6, 19

+35

32-38; oe 13-22; C, 19

3248: B, 13-18

, 2, 35; B, 0-7, 22; C, 2-19: Masillon I. in

o>>>

32-38, 55; B, 13-22

2, 35; B, 6, 13-18; C, 6-19

o> > >>

"92: ion 19-20

A, 32-38: B, 22; C, 6-19

B, 13-18

A, 32-38

A, 10, 32-38; B, 13-22; C, 19

A, 13, 32-38; B, 6-22; C, 10-19

A, 13, 32-38

A, 32-38

C, 10-18, on Cystephora brownii

C, 19-22: D, 3, 4, on Posidenia australiy and

P. osienfeldit

*, 6

, 6, 19-20: D, 3, 4. on Posidonia australis

and P. ostenfeldii

A, 32-38; B, 13-18

A, 32-38

D, 3, on Posidonia australis

C, 19-20

A, 5-38: B, 5-32; C, 8-20

A, 6-38; B, 4-32; C, 8-19, Masillon L. in

bay, 14

A, 0-2;,C, 0-3

SUBTIDAL ALGAL AND SEAGRASS ECOLOGY 189

Cystophora gracilis Womersley & Nizamuddin

Cystophora monilifera J. Agardh

Cystophora moniliformis (Esper) Womersley &

Nizamuddin

Cystophora pectinata (Grev. & C.Ag.) J. Agardh

Cystophora subfarcinara (Mert.) J. Agardh

Myriodesma harveyanum Nizamuddin & Womersley

Sargassaceae

Phylotrichia

Sargassum decipiens (R.Br.) J. Agardh

Sargassum heteromorphum J. Agatdh

Sargassum varians Sonder

Sargassum verruculosum (Mert.) Agardh

Arthrophycus

Sargassum bracteolosum J. Agardh

Sargassum lacerifolium (Turn,) Agardh?

Sargassum trisfichum Grey. & Agardh ex Sonder

Eusargassum

Sargassum linearifoliam (Turn.) Agardh?

Sargassum podacanthum Sonder?

Sargassum spinuligerum Sondet

Sargassum distichum Sonder

Sargassum (Eusargassuin, tribe Glomerulatae?)

RHODOPHYTA

Nemaliales—Chaetangiaceae

Galaxaura spathulata Kjellman

Helminthocladiaceae

Liagora harveyiana Zeh

Bonnemaisoniaceae

Asparagopsis armata Harvey

Delisea hypneoides Harvey

Delisea pulchra (Grey.) Montagne

Gelidiales—Gelidiaceae

Pteracladia lucida (R.Br.) I. Agardh

Cryptonemiales—Dumontiaceae

Acrosymphyton taylori Abbott

Squamariaceae.

Senderephycus australis (Sond.) Denizot

Corallinaceae (excluding encrusting taxa)

Amphiroa anceps (Lamarck) Decaisne

Jania fastigiata Harvey

Jania micrarthrodia Lamouroux?

Jania pusilla (Sond.) Yendo

Jania sp,

Corallina cuvieri Lamouroux

Corallina cuvieri f. crispata Lamouroux

Metagoniolithon charotdes (Lamx.) W. v. Bosse

Metagoniolithon stellifera (Lamarck) W. y, Bosse

Poly porolithen patena (H. & H.) Mason

Cryptonemiaceae

Carpopeltis phyllophora (H. & H,) Schmitz

Cryptonemia undulata Sonder

Halymeénia harveyana ¥. Agardh

Thamnoclonium dichotomum (J.Ag.) J. Agardh?

Grateloupiaceae

Gelinaria ulvoidea Sonder

Kallymeniaceae

Austrophyllis alcicarnis (J.Ag.) Womersley & Norris

Callophyllis rangiferinus (Turn.) Womersley

Callophyliis lambertii (Turn.) J, Agardh

Kallymenia cribrosa Harvey

Kallymenia spinosa Womersley & Norris

Thamnophyllis lacerata Womersley & Norris

Gigartinales—Plocamiaceae

Plocamiumn angustum (3.Ag.) Hooker & Harvey

B, 4-7

A, 0-2; C, 2-19

B, +7

A, 6, 13; B, 6-18; C, 10-19

B, 6; C, 0-13, 19, Masillon I. in bay, 1-4

A, 2; B, 4-7

. 32-38; B, 6-18; C, 6-19

» 35; B, 13-18; C, 2-19

A, 13, 32-38; B, 4-22; C, 10-19; D, 2

A, 12, 32-38

Masillon I. in bay, 1-4

A, 32-38; B, 22. Masillon I. in bay, 1-4

A,2

A, 10, 32-38; B, 4-18

A, 32-38; B, 13-18; C, 19

A, 10, 32~39;,B, 6-22

B, 4-7, Masillon J. in bay, 1+

A, 32-38

B, 13-32

A, 32-38; B, 13-18, 32; C, 10-19

C, 0-2

D, 3 on Postdonia ausiralis

B, 4-7; Masillon I. in bay 1-4 on Cysto-

phora subfarcinata

A, 32-38

C, 6-1

D, 2 on Amphibolis antarctica

B, 13-18 on Ballia caltitricha

A, 32-38, Masillon I. in bay, 1-3

D,2

B, 13-18; C, 10-13

A, 32-38

C, 19-20

A, 32-38; B, 13-18, 32

A, 2-10; B, 0-7; C, 10-13

A, 55

A, 32-38: B, 13-22

A. 55

‘A. 32-38

A, 6-55; B, 6-32; C, 6-19

190 S. A. SHEPHERD & H. B. 8S. WOMERSLEY

Plocamium cartilagineum (.) Dixon

Placamium leptophylium Kuetzing

Plocamium mertensii (Grev.) J. Agardh

Plocamium preissianum Sonder

A, 6—10, 32-38, 55; B, 13-22, 32

A, 55

A, 6-55: B, 6-22: C, 6-19

A, 6-55: B, 6-32; C, 6-19

Solieriaceae

Solieria robusta (Grev.) Kylin A, 32-38; C, 20; D, 4

Rhabdoniaceae

Areschougia congesta (Turn.) J. Agardh? A, 32-38

Rhodophy!lidaceae

Rhodophyllis membranacea (H. & H,) Harvey

Rhodophyllis ramentacea (C.Ag,) J, Agardh

A, 35: B, 13-22

A, 32-38; B, 32

Hypneaceae

Aypnea episcopalis Hooker & Harvey B, 6; C, 10-13

Alypnea sp. A, 2; B, 46

Mychodeaceae

Mychodea pusilla (Harv.) J. Agardh D, 2, on Amphibolis antarctica

Mychodea ramulosa J. Agardh B, 4-7

Mychodea carnasa Hooker & Harvey A, 32-38

Neurophyllis australis Zanardini C, 19-20

Dicranemaceae

Dicranema revolutum (C.Ag.) J,-Agatdh

Acrotylaceae

Hennedya crispa Harvey

Rhodymeniales—Rhodymeniaceae

Fauchea?

Webervanbasyea kaliformis (J.Ag.) J. de Toni

Webervanbossea splachnoides (Harvey) J. de Toni

D, 2, on Amphibolis antarctica

D, 24, loose-lying

A, 32-38

A, 32-38; B, 22; C, 10-13, 19

A, 32-38: C, 19-20

Botryocladia obovata (Sonder) Kylin C, 19-20

Coelarthrum cliftonit (Harv.) Kylin A, 32-38

Caelarthrum meulleri (Sond.) Boergesen A, 35

Gloiosaccion brownii Harvey

B, 13-18; C, 10-13

Rhadymenia australis (Sond.) Harvey

A, 32-38, 55; D, 2

Lomentariaceae

Champia affinis (H, & H.) J. Agardh A, 32-38; B, 13-18; C, 19

Champia obsoleta Harvey A, 2

Champia tasmanica Harvey A, 32-38

Ceramiales—Ceramiaceae

Crouani¢ae

Gartya pinella Harvey A, 55

Gulsonia annulata Harvey C, 19-20

Antithamnieae

Acrothamnion preissii (Sond.) Wollaston B, 13-18, on Ballia eallitricha

Antithamnion divergens (J,Ag.) J. Agardh A, 35

Ballia ballioides (Sond.) Wollaston

Rallia callitricha (Ag.) Kuetzing A. 33 B, 13-18, 32

A, 55

Ballia mariana Harvey 3

Platythamnion nov. sp? A,55

Griffithsieae

Griffithsia tezges Harvey A,2

Callithamnieae

Callithamnion sp. A, 32-38

Callithamnian sp. A, 55

Dasyphileae

Rhodacallis eleguns Kuetzing A, 55

Delesseriaceae

Apoglossum tasmanicum (F.v.M.) J. Agardh B, 32

Dasyaceae

Dasya clavigera (Wom.) Parsons C, 0-2; D, 24

Dasya naccarivides Harvey? C, 19-20

Rhodomelaceae—Polysiphonieae

Poalysiphonia nigrita Sonder A,2

Pterosiphonicae

Pterosiphonia sp. B, 6

Herposiphonieae

Dipterosiphoniu? nov- sp? B, 13-18

Herposiphonia nov. sp? A, 55

SUBTIDAL ALGAL AND SEAGRASS ECOLOGY 191

Polyzonieae

Clijtenaea pectinala Harvey

Amansieae

Osmundaria prolifera Lamouroux

Laurencieae

Lawurencia elotu (Ag,) Harvey

Laurencia filiformis (Ag.) Montagne f. dendritica

Saito. & Womersley

A, 32-38; B, 13-18; C, 19

B, 13-18; C. 6-20

B, 13-18

A. 32-38

Laurencia filifarmis (Ag.) Montagne f. heteraclada

Saito & Womersley

Laurencia paniculata (Ag.) J. Agatdh

SPERMATOPHY TA—seagrasses

Potamogetonaccac

Heferozostera tasmanica (Mart. ex Aschers)

den Hartog.

Posidonia australis J. D. Hooker—narrow. and

broad forms

Posidonia ostenfeldti den Hartog

A, 0-2; C, 19-20

C, 19-20

D, 3

C, 19-22: D, 3-4

D, 3, Masillon I, in bay 5-9

Amphibolis antarctica (Labill.) Sonders ex Aschers D,2

Acknowledgments

We are gtateful to the Royal Society of

South Australia (Research and Endowment

Fund) and the Department of Fisheries for

grants towards the field work. The first author

was assisted in the SCUBA work by Mr K. L.,

Branden and Mr N, Coleman, to whom thanks

ate expressed. Assistance from the Australian

Research Grants Committee in provision of the

technical assistance of Mrs E, L. Robertson

and Miss C. Anderson, is gratefully acknow-

ledged by the second author.

References

Boupouresgur, C. F, (1971).—Contribution a

"Etude Phytosociologique des peuplements

alzaux des Cotes varoises. Vepetatio 22, 83-

184,

CHapman, V. J. (1967).—The sublittoral fringe in

the Pacific. J. Indian Bot, Soc. 46, 337-343.

CSLR.0, (1967a)—Aust. Oceanogr. Cruise

Report No. 34.

CSTR.O. (1967b)—Aust. Oceanogr. Cruise:

Report No. 46.

C.S,LR.0, (1968)—Aust. Oceanogr. Cruise

Report No. 43.

C.S.LR.0. (1969).—Aust. Oceanogr, Crulse

Report No, 54.

Gris, R. W, & Kwara, R. S. (1970).—Some

ecological effects of discharged wasies on

maziue life. Calif. Fish and Game 56, 145-

185.

JERLOV, N. G. (1968).—"Optical Oceanography”.

(Elsevier: Amsterdam.)

Lining, K, (1970).—Tauchuntersuchungen zur

Vertikalverteilung der sublitoralen Helgoliin-

der Algenvegetation. Helgoldnder — wiss.

Meeresunters 21, 271-291.

Mann, K. ( 1972), —Ecological energetics of the

seaweed zone in a marine bay on the Atlantic

coast of Canada. |. Zonation and biomass of

seaweeds. Mar. Biol. 12, 1-10,

Rrept, R. (1971)—"“Water movement”. In 0,

Kinne (Ed.). “Marine Ecology. A comprehen-

sive integrated treatise on life in oceans and

coastal waters”. Vol. 1, Part 2, pp. 1123-1156,

(Wiley—Interscience: London.)

Rrepr, R. & Forsrner, H. (1968).—Wasserbewe-

gung B Mikrobeyeich des Benthos. Sarsia 34,

163+!

SHEPHERD, S. A, & Womerstey, H. B. S.

(1970).—The sublittoral ecology of West I.

South Australia. 1, Environmental features

and the algal ecology, Trans. R. Sov. §. Aust.

94, 105-138.

SHEPHERD, S, A. & Womensiey, H. B. S.

(1971) —Pearson Island expedition 1969, 7.

The subtidal ecology of benthic algae. Trans.

R. Soc. §. Aust. 95, 155-167.

Stracuan, A. R. & Kosxr, R. T. (1969) —A

survey of algue off Palos Verdes Point, Cali-

fornia, Caltf, Fish and Game 55, 47-52.

Vaux, D, (1970).—Surface temperature and

salinity for Australian waters 1961-65.

Si ‘S.LR.O. Aust. Div. Fish. Oceanog. Atlas

o. 1.

WHittagker, R. H. (1967),—Gradient analysis of

vegetation, Biol, Rev. 42, 207-264.

Womras.ey, H. B.S. & Epmonps, 8. J. (1952).—

Marine coastal zonation in southern Australia

in relation to a general scheme of classifica-

tion. J. Ecol. 40, ‘84-90.

Womersiey, H. B. S. & Epmonos, 8. J. (1958).—

A general account of the intertidal ecology of

South Australian Coasts. Aust, J. mar. freshw,

Res. 9, 217-260,

BREEDING BIOLOGY AND LARVAL DEVELOPMENT OF

LITORIA VERREAUXI (ANURA: HYLIDAE)

BY MARION ANSTIS**

Summary

ANSTIS, M. (1976).-Breeding biology and larval development of Litoria verreauxi (Anura:

Hylidae). Trans. R. Soc. S. Aust. 100(4), 193-202, 30 November, 1976.

Oviposition and larval development of the hylid frog Litoria verreauxi are described and ecological

notes are given. Comparisons are made with other hylid frogs (particularly members of the Litoria

ewingi complex).

BREEDING BIOLOGY AND LARVAL DEVELOPMENT OF

LITORIA VERREAUXI (ANURA: HYLIDAE)

by Marton ANSTIS*

Summary

Awnstis, M, (1976),—Breeding biology and larval development of Litoria verreauxi (Anura:

Hylidae). Trans. R. Soc. §. Aust. 100(4), 193-202, 30 November, 1976.

Oviposition and larval development of the hylid frog Litoria verreauxi ure described and

ecological notes are given. Comparisons are: made with other hylid frogs (particularly members

of the Litoria ewingi complex)...

Introduction

Litoria verreauxi (Duméril), previously in-

cluded in Hyla ewingi Duméril & Bibron (see

Littlejohn 1963, 1965; Tyler 1971) is a hylid

frog found along the coast of eastern Australia

from Victoria to southern Queensland (Little-

john 1965; Straughan 1966)+. Adult morpho-

logy in the Sydney areca has been described by

Copland (1957) as H. ewingi verreauxi, and

by Moore (1961} as HA. ewingi. Fletcher

(1889) and Harnsen (1922) provided some

data on the breeding season, ova and larvae,

while Moore (1961) briefly described advanced

embryos and larvae. Martin (1965) described

tadpoles from the Melbourne area bul did not

diseuss embryonic development. Martin &

Watson (1971) mention some life history

characteristics. The present paper provides data

on breeding biology and larval ecology and

includes a detailed description of embryos and

larvac.

L, verreauxi appeat'’s to be related to a com-

plex of species including L. ewing?, L. parae-

wingi, G. jervisiensis, and possibly Z. Burrawsl

(Martin & Littlejohn 1966; Martin 1967a;

Watson, Loftus-Hills & Littlejohn 1971).

Where data arc available, comparisons ure

made with these taxa.

Material

Six egg masses of ZL, verreanxi laid in the

laboratory, together with samples of larval

material from the field, form dhe basis of the

study. Egg masses came from an adult popu-

lation, originally collected at Darke’s Forest in

1970 and released in a garden at Penshurst.

Frogs from adjacent areas in Penshurst may

also have joined the population.

An egg mass from a pair of ZL. ewingi cap-

tured in amplexus at Lobethal, 5. Aust. on

30.vili.1972, was maintained to hatching stages.

Larvae of L. paraewingi from 2 km N of Glen-

burn, Vict. were examined for comparison.

Collecting loculitics and dates are listed in

Table 1.

Methods

A series of outdoor aquaria containing rain-

water and vegetation was maintained at Pens-

hurst and checked regularly for the presence

of spawn. Three pairs (one in amplexus) were

captured in the vicinity of the aquaria (two on

11.ix.1972 and one on 20.11.1974) and placed

in plastic bags containing water, twigs and

vegetation, Oviposition behaviour of these three

pairs was studied.

Embryos were maintained up to stage 25 in

shallow water ranging from 14°-21°C. Larvae

from the various localities were maintained

separately in open outdoor aquaria, and indi-

viduals from some were reared to metamor-

phosis. The behaviour of laryac was studied

both in aquaria and at field collecting sites.

Food provided consisted of algac and other

water plants, commercial fish food, boiled let-

tuce and occasionally meat. Water temperature

during larval development ranged 8°—27°C.

Specimens from each group were fixed at inter-

* 630 King George’s Road, Penshurst, N.S.W, 2222.

t Straughan, I. R. {1966)—An analysis of species recognition and species isolation in certain Queens-

lattd frogs. Ph.D. thesis, University of Queensland (t:npuhl.).

194 MARION ANSTIS

TABLE 1

Breeding sites of Litona verreauxi

Collecting

Locahty Description of habitat date Stages Other Iatvae present

Menai, 1, Permanent dam in dry sclerophyll bush- 21.11.1971 34-42 Liroria aurea

34°02'S land. Surface vegetation, rooted plants, L, latopalmata

151°01" EB mud substratum. Uperoleia marmoraja

Ranidella signifera

2, Concrete water vessel, permanent water, 16.ix.1972 34-41

surface vegetation, mud substratum

Penshurst, Permanent ouvidoor aquaria jn suburban gar- Numerous 1-46

33°58'S den, Surface and rooted plants dates, 1970

151°05' E to 1974

Darke’s Forest, 1, Permanent flowing stream, sandstone base, 16.ix.1972 16-18 Liforia jervisiensis

34°12'S fast. flowing sections, deep pocls in dry

151°S8"R sclerophyll bushland,

2. Permanent dams, liitle rooted and no sur- 24.in.1972 26-40 Limnodynastes peroni

face vegetation, mud substratum 30. x.1972 Litoria perori

2,x1,1972 Ranidella signifera

6.A7.1972

Qurimboh, Semi-permanent, small, slowly flowing creek, 9.in1973 25-33 Ruanidella sienifera

33°22°S shallow pools, rooted vegetation, mud sub-

151°22"E stratum. Cleared farmland in wét Sclerophyll

forest

Glen Alice, Semi-permanent, shallow pond, grass but- 1vi.l974 25-28 Limnodynastes

33°02’ 5 tom, in open cleared farmland with sur- tasnaniensis

151°12’E rounding woodland

‘Spring Creek, Permanent creck, slowly fowing small 25,1.1973 30-46 Mixephyes balbus

30°29°S§ pools, sandy and basalt substratum. Wet 25.07.1973 25-42 Ranidella signifera

152°24" B sclerophyll forest, paruy cleared 18.iv.1973 Litoria glandulosa

L, pearsoni

Dorrigo, Small, slowly flowing creek, surface vegeta- 26.41.1974 28-43 Mixophyes fasciolatus

30°20 S tion, mud substratum. Cleared rainforest Adelotuy brevis

182°43°E farmland

Rouse Hill, Permanent waterhole in cleated paddock, 19.x311972 27-42 = Liloviavaeruiva

33°42'S Dry sclerophyll bushland area, farmland Ranidella signifera

150°55°E

Fig. 1. Lateral and dorsal views of farya showing

measurements for morphometric charac-

ters,

vals in 4% formalin, after being relaxed in 1%

chlorbutol solution; larger specimens were

injected with a small quantity of formalin

before final fixation.

Measurements were taken with verier cal-

lipers reading to 0.1 mm or an ocular micto-

meter (reading to 0.01 mm). Drawings were

made using a drawing tube attached to a

stereoscopic microscope. All measuzements and

drawings are based on preserved specimens,

while descriptions are of both preserved and

live material. The staging system used is that of

Gosner (1960). Abbreviations and definitions

of latval morphometric characters (Fig. 1)

are: ST—total length (tip of snout to tip of

tail); BL—body length (tip of snout to junc-

tion of body wall and tail musculature); BW—

maximum body width; BD—maximum body

depth; TD—maximum tail depth; TM—depth

of tail musculature (measured in line with

TD): 10—inter-orbital span (minimum dis-

tance between the cyes, measured at the central

inner edge of each eye); [N—internarial spazt

BREEDING BIOLOGY OF LITORIA VERREAUXI 19s

(minimum destance from eye to avis); EN—

distance from eye to narts; MW—maximum

width of oral disc,

Results

Calling aetiviry: The mating call has been des-

cribed by Littlejohn (1965), Males alt Pens.

hurst call throughout the year, with the most

intense activity on mild, wet nights during

spring and summer. Diurnal calling mostly

occurs during and after rain. Males call while

afloat near the edge of ponds by night, or from

low vegetation or ground near the water by

night or day. At 2300 hrs on 20.11.1974 at

Penshurst, during light rain, a silent male sur-

faced in an aquarium gbout 4 cm from a cuall-

ing male. The latter turncd to face the former

and, after a brief pause, swam slowly towards

him, calling in softer, separate totes (quite dis-

tinct [rom the mating call) and attempted

amplexus. The silent male immediately swam

of, The calling male did not follow, but

resumed a normal mating call.

A similar behavioural sequence preceded

ampleaus in one of the pairs captured on

11.ix.1972, the male emitting soft. separate

notes as he approached the female,

Ovipesition: Oviposition at Penshurst has been

observed in February, March, June and Sep-

tember—December, The following description is

a composite of observations of the three pairs

studied.

When frogs were collected on 11.x.1972, air

teniperatures 2 cm above water were {8°-]9°C

and surface water temperatures 19°—23°C.

Amplexus commenced in these pairs at 2000

and 2325 hrs, Eggs were laid in scparate

hatches attached to twigs or reeds over a period

of hours (Tuble 2). Before oviposition, the

female showed lateral abdominul contractions,

either simultancously or alternately. These con-

traciions usually became more powerful as ovi-

position was near and lasted about one second,

with two or more occurring In succession.

In a typical behavioural sequence, a pair

submerged and the female grasped a twig with

one hand. She dorsiflexed her body with the

hind limbs extended and, as the batch emerged,

Fig. 2a. Oviposition with the miale receiving, und

fertilising the eggs.

Fig. 2b, The mate pushes the batch down to the

female's feet.

the male lowered his vent towards the eggs

and cupped his feed areund, so holding them

(Fig. 2a). The sides of the male then undu-

fated und his feet moved up and down in a

bref fanning motion over the eggs. This pro-

cess of oviposition and fertilisation lasted 3 sec,

The female ventriflexed, drawing her legs back

under her hody, and the male rolled the batch

down to her feet (Fig 2b). The female held

the hatch motionless for 40 sec. She then

pulled herself around the twig in spiral fashion,

wrapping the eggs round it with her fect. The

pair left the eges and returned to the surface.

After 1.5-7.3 min. the entire process was

repeated, and S min.—2 he clapsed before fur-

ther batches were laid.

Variations were: (1) Nearing the end of

amplexus, two or three batches were laid in

very close succession, each being held by the

feet of the feinale for 40-60 sec. before the

ensuing one was laid, The resulting composite

batch was then attached to supporting material.

(2) Females varied in their attempts to spread

TABLE 2

Ovipovition bohtavionur

Duration of

‘Total cyiragion Dirratian at ege- single batch Batch holding, Total eges

Paic of Ainplexas laying period oviposition time (female) n laid

Ta 5 hr 15 min, 2 hr 8 min. Tt j0, 35-60 sec, 15 757

Z Unknown 3 hr 47 min. 2-4 sec. 35-60 sec. Bors 101i

196

batch one

laid and held

by femate for

40 seconds

single batch

laid and held

by female for

40 seconds

batch 2 laid,

batches 142

held together

for 40 seconds

eggs attached

to vegetation,

adults return

to surface

batch 3 faid,

batches 1,243

held together

for 40 seconds

Fig. 3. Oviposition cycles during which @ single

batch is tald and attached to vegetation,

or two or three are Juid in close succes-

sion before attachment,

ott the eggs in spiral fashion, sometiines

swivelling around the twig only once or not at

all, resulting in thicker clumps of eggs. (3)

One female used her left hand to grasp and

pull free some eggs which had adhered to her

venter, before attaching the batch to vegeta-

tion,

Females had more difficulty in wrapping a

composite batch around a twig, and offen aban-

doned the egexs as a thick mass. In aquaria

lacking vegetation or twigs, egg masses have

been found in thick clumps on the substratum,

in water up to 50. cm deep,

The final stages of armplexus in one pair

were: at 0100 hrs the female made move-

ments similar to the croaking motions of males,

but produced no sound, At O108 hrs she

submerged and both male and female began

typical ovipositional behaviour, but the female

remained in the dorsiflexed position for 7.7 sec,

(4.7 sec, longer than average) and produced

no eggs. Two sec, Jater the pair fell apari, both

floating motionless on their sides just under

the surface, with limbs tightly adpressed

against the body. After LO sec, the male

MARION ANSTIS

Fig. 4, Two batches of eves joined and attached

io a stem, Filamentous algae are catwined

amongst the egg mass.

recovered from this state of suspension and

surfaced, the female doing so 5 sec later. A

second pair behaved similarly, except that the

period of motionless suspension was shorter.

The basic cycle of oviposition behaviour is

shown in Fig. 3. Laying of all eggs conyprises

a number of such cycles.

Qva: In natural environments egg masses are

altached to submerged reeds, twigs or grasses

usually close to the surface (Littlejohn 1963),

The cgys cohcre and the inner ones stick ta

the supporting material. There is a single layer

of jelly around each egg, but within « roass the

individual capsules merge and are not clearly

defined (Fig. 4).

The mean diameters of eggs and capsules in

stages 1 and § are shown in ‘Table 3. Ova

generally have a dark brown animal pole and

an off-white, yellow or orange Vegetal pole, All

ova from a single female arc the same colour.

The animal pole gradually lightens from gas-

trulation onwards.

The number of eggs in 20 single batches

ranged from 1—52 (mean 30). Three “double”

batches contained 64, 78 and 79. The totul

complements of four females were 1,011, 757,

632 and 522.

Development of embryos; After Fertilisation

there is no distince grey crescent. Cell division

appears normal, although not as symmetrical

as in Gosner’s (1960) diagrams. The vegetul

pole always divides later than the animal pole.

At stage 17 (tail bud: Fig. 5a), the head

region is well defined, showing optic bulges, gill

plates, U-shaped adhesive organ and a slight

stamudaeal pit. The posterior crescent of the

adhesive organ is less distinct. Tn some embryos

BREEDING BIOLOGY OF LITORIA VERREAUNI 197

the visceral arches and a slight pronephric

bulge are discernable. The tail bud is straight

anu points dorsally, with no obvious tail fin

rudiment, In late stage 17, just before muscular

movement begins, the tail bud extends and

points cither fo the right or to the left, and the

posterior crescent of the adhesive organ almost

disappears, yielding two separate organs which

ute heavily pigmented. Embryos in stages 17

to 20 have a yellow yolk sac and are light

brown elsewhere.

The embryos begin hatching when. they have

reached stages 19 and 20, At stage 20 (Fig, 5b)

the gills are small, just functional and non-

pigmented. The optic bulges are more defined,

and there is a small crescent of melanophores

around the anterior edge of each, The stomo-

dacal pit bas deepened and the adhesive organs

are prominent. The yolk sac has elongated and

18 wenerally narrow, and there are small areas

of pigment along its dorsal edge, and between

the optic bulge and olfactory pit, The arca

above the olfactury pit. fy clearing and the tail

fing aré a translucent milky white.

With the temperature regime prevailing

during carly development, hatching was com-

plete after 147 hr when most embryos were

in stages 21-23. The external gills are fully

developed in stage 2] (Fig. 5c). The tail fins

and cornea clear during stage 22; the oper-

culum partly covers the gills, and the distribu-

tion of melanophores increases over the yolk

sac, beneath the eyes, around the nares and

along the dorsal surface of the tail muscula-

ture. At stage 23 the gills are reduced, the

external nares are open, the stomodaeal pit

deepens further and the oesophagus begins to

differentiate. The anal tube is developing and

the fins, how transparent, take on their charac-

teristic arched shape. Generally, pigrnentation

increases, dispersing into the pattern typical of

the larva. The yolk sac is pale yellow beneath

the layer of melanophores, while other dorsal

and fateral areas surrounding the pigment, be-

come transparent. However one group of em-

bryos at this stage lacked dark pigment (except

for the eyes), and appeared yellow. These

embryos did not develop melanophores until

stage 25,

At stage 24 the mouth-parts have developed

oral ridges and a stnall non-keratinised beak,

the oral suckers have diminished, and the oper-

culum Closes on the right side. The anal tube

is partly open in some cmbryos, During stage

25 the formation of mouthparts is virtually

completed, the beak becoming keratinised and

TABLE 3

Dimensions in.mm of embryos and larvae of L.

verreauxi from Penshurst

(meuns, with canges in brackets)

Embryos

Embryo Capsule

Stege nh diam. diam.

i 10 i.23 43

{1.19-1.23) (3.53-4.92)

# 8 1,20 4.55

(1,15-1,23) (4.26-4.92)

9210 9 1.28 3.88

(1.23-1.39) (3.44 4.35)

14 9 1,58 4.28

(1.48-1,64) (3.74-4.929

15 y 4.74 441

(1.68) 1.80) (4.10-4,92)

7 1m Zt 4,39

(1,.85—2.4)) {3.61-6,40)

Embryea

Stare a diam.

20 10 5.83

(5,62-5.991

21 10 6,27

(6.76.44)

22 tt 640

(6.15-6.64)

23 id 6.98

(6.23-7.30)

24 9 7.26

(6.72-7.71)

25 it B45

(7.87-9,18)

Larvae

Stage n Body length Totablength

26 ju 10,16 33.6

(9.02-12,79) O83 ba)

27 10 Vth 24,2

(10:50-13,64) (21,2-27.2)

13 10 10.85 24,0

(9,68-11,91) (214-27,6)

a 9 114.16 24,5

(10.33-33,97) (24.4-272)

3b 10 12.88 29.)

CUE, U5-39,.78) (25.2-31.5)

3h 10 13,65 334

(12,30-15,42) €27,5-39.4)

2 7 13.40 305

(11,91-14.27T) (27.0-33,2)

WY s 14.31 32.6

{19.94-15.58) (31,0-44.0)

4 & 14,85 34.7

(13.02-15,74) (30,1-37.6)

Ws 16 16,65 4l.i

£14.76-19.68) (33.0-48.8)

46 10 16,15 41.4

(15.35-13.61) (34,447.23

a7 3 16.22 39,5

(14,92-17,22) (36.4-44.8)

198 MARION ANSTIS

Stage fn Bodylength Totallengih

38 6 16.84 432

(15.00-18.00) (39.6-46.0)

34 6 17.27 45.6

(16:73-18,32) — (42.0-51,9)

a it 16.87 46.6

(14,76-18,37) (39,2-52,2)

ay i 17.09 48,2

(16,56-18,20) (45,0-52.9)

4 3 16,13 43.1

(14.27.1738) (40.5-45,1)

a3 4 14.54 34.6

(14,.27-14,92) —-137.4-39.7)

45 1 16.3 —

(14,9-18.6)

46 la 15.3 =>

{13,.2-17.5)

lablal teeth developing on the oral ridges. The

labial papillae may not reach their total num-

ber until stage 26 or later. The spiracle be-

comes functional und the anal tube is fully

open. The remnant adhesive organs praduully

disappear during this stage.

Measurements of embryos are shown in

Table 3,

Larvee: A composite description of 10 larvae

at stage 35 (Figs 5d-f) from Penshurst fol-

Jows; Body widest across the mid region of the

abdomen and ovoid. Snout evenly rounded fn

dorsal view and tapers to a truncate edge in

lateral view.. Nates dorsal and raised on very

short tubes which open antero-laterally. Eyes

lateral and relatively large, Spiracle sinistral,

ventrolateral and not visible from above, Tt

opens in a porstero-dorsal direction and dia-

meter of the spiraculat tube decreases slightly

from its origin 10 its opening, Anal tube dex-

tral, very short, of small diameter and opens

about halfway up the ventral fin. Tail fins

arched and taper to a fine point. Dorsal fin

extends midway up the body, deepest approxi-

mately halfway along its length, Ventral fin

deepest along its anterior third. Tail musoula-

ture moderately thick, narrowing to a fine point

posteriorly.

Mouth antero-ventral in position and has

border of papillae around all but the anterior

margin (Fig. 6). In some specimens. there is

also a median gap along the posterior murgin

(possibly caused by damage). Papillae most

numerous laterally. Two upper and three lower

rows of Jabial teeth, two upper being of

approximately equal length in most specimens,

First two rows in the lower fabiim are also

about equal, third lower row is usually the

Fig, 5. Embryological and larval development of

Litoria verreauxi, Penshurst, (Bar rep.

resents 1 mm). Stages: 2—i7, b—20, c—

21, d—36, e—1h, f—46.

shortest. In same specimens a partial median

gap occurs in secand lower row and other rows

may be interrupted at various points, probably

through damage. Beaks of moderate propor-

tions, serrations fine on inner edge of lower

beak and very fine on the upper beuk.

The only consistent geographic variation

noted Was In specimens from Spring Creck,

most of which had more massive beaks and

two pigmented areas below the lower beak

(Fig. 6b). Specimens from Dorrigo also

showed a tendency towards more massive

beaks. It was noted that specimens from the

northern localities generally had shallower fins

than most southern specimens (Table 4), Body

dimensions of larvae are given in ‘Table 3,

In hfe the dorsal surface varics amongst

individuals from light golden to a very dark

BREEDING BIOLOGY OF LITORIA VERREACXI 199

Fig. 6. Mouthparts of Z. verreauxt. a, {tom the

southern site cf Penshurst; bh, from the

northern site of Spring Creek (bar rep-

resents | mm).

brown. (almost black). In some specimens the

pigment ts motiled, The areas of skin over the

trabeculae cormba, central nervous system

{brain und spinal cord to base of tail). the

abdumern ond surrounding the nares, are

darker, There is a copper-gold sheen ventrally

and laterally over the abdomen. In lateral view

the areas covering the pharynx and buccal

cavity (excluding eyes) are transparent (except

for some melanophores héetween the eye and

mauris}, and the pills, heart and developing fore-

timbs are visible. From the ventral aspect the

areas over the gills, heart and buccal cavity are

unpigmented.

The tail musculature ig cream with irregular

dark blotches over the dorsal surface, and

partly over the lateral surface. In generally

darker larvae the musculnture may be uni-

formly pigmented. The dorsal and ventral fins

TABLE 4

Propartions.in mar af L. verreauxi larvae from

different localities

(means, with ranges im brackets)

Narthern Southern

(Spring Creek, Dolnee) ‘Penshurst)

Stage 33 & Sh 45 &36

q 7 10

ST 34.1 431

(3u9412) (36.2-48.8)

BL 11.69 17,27

412,4G-18.56) (18,53-19,68)

BW 774 1u.26

(6.72-R.36) (8,69.717.48)

BD 779 W507

16,56-4.95) (8.86-12,14)

TD TSB 119

16.48.63) (8.59- 7230)

mM 2.68 3.60

€2.13-3.28) (2,79-4,59)

{Q 3,62 5.53

€3.44-4.55) (4.66 6.40)

IN 2.05 2.64

(1.72-2.21) (2.46-2,95)

EN 228 2.65

(2.13-2.69) (2.38-3,12]

MW 2.03 4.10

(2,95=4 43) (3,6]-4.66)

wary from dusky (in dark larvae) to aimost

transparent (lighter larvac), with parts of the

tail vascular system pigmented, Larvae with

mottled pigmentation over the body also have

mottled tails. The iris. is golden.

Specimens which were dark in life may

fetain much of this pigment in preservative,

Those which were light golden hecome an off-

white colour in all but the darker areas. and

the skin is clearer than in fife. The copper-gold

sheen is lost and the abdomen may appear

dark shiny blue fur some time in preservative,

then eventually turn black. The iris loses its

golden colour and also appears: black.

Lurval hehaviewr> After hatching the embryos

remain close to the egg capsules until about

stage 24. Durning stages 25 to about 27, the

larvac are most often found in the shallow

areas of ponds, particularly near the edge, but

beyond this stage a much greater water space

is utilized.

The larvae are of the active, nektonic type

{Orton 1953) and spend much of their time

hovering in the water by rapidly oscillating the

Jail tip (flagellum). They frequently cruise

slowly to the surface with head uppermost jt

ahout a 45° angle, using only the flagellum for

propulsion. When feeding at ihe surface, they

often position themselves almost vertically and

can remain suspended at this, or any level in

the water. They arc capable of sudden spurla

of speed (during which they may use the entire

tall and body), and rapid changes of direction

(making use of the deep fins), when disturbed,

As well as feeding at the surface, the larvae

Eruge on vegetation and other material in any

zone of the pond and scavenge: in bottom sedi-

ments. The variation in larval pigmentation

appears to be related to characteristics of the

habitat, Specimens in muddy water, or clear

water over a dark substratum, usually range

from dusky brown to almost black, while

those in clear water aver a light snbstratum

tend to be golden, with the darker areas con-

trasting, but less pronounced,

Larvel life span and meramerphosis; Metamor-

phosis of larvae reared from cges laid sat

Penshurst on 11.ix.1972 began on 10.xti.1972,

giving a spring-summer larval life span of 90

days. Metamorphosis of larvae from egg

masses laid on 23,%,1971 occurred from Jate

December to early March. Metamorphosis was

also recorded at Penshurst fn September 1972

and at Menai from 27-29.iK.1972, It is there-

fore known to occur from September—March,

but probably takes place at other times because

ege masses haye been found jn most months

nf the year.

The body fengths of 10 juveniles at stage

45. and 18 at stage 46 are shown in Table 3.

Al these stages the juveniles closely resemble

the adults in colour, but lack the deep orange

of the anterior and posterior surfaces of the

thigh, and the black spots in the groin. Pale

orange thigh colouration is visible in some

juveniles at stage 46,

Discussion.

Calling aciivity: Fletcher (1889) and Harrison

(1922) moted that calling occurs throughout

the year, and Moore (1961) observed calling

activity from the end of July 1992 to late April

1953. Watson ef al (1971) record calling acti-

vity in all months except July and found that

1, verreauxi males when sympatric with L.

ewitel usually call on Jand up to 25 m from

water, and only rarely in water. This latter be-

haviour contrasts with that of males at Pens-

hurst and Darke’s Forest which commonly call

in water.

A call distinct from the mating call, given by

the male on approaching a potential rival or

mate, has been observed; its function is not

known, More abservalions aré necessary to

MARION ANSTIS

establish the extent of behavioural variation iu

this species. A similar call has been observed

in L, ewfng! (Anstis 1976).

Oviposition: Harrison (1922) found spawn in

Sydney every month of the year, and Moore

(1961) collected embryos in August, 1952 at

Killafa. Fletcher (1889) found a pair in

amplexus in June, 1885 and stated that the

species “probahly breeds nearly throughout the

year", This agrees with the oviposition dates

recorded at Penshurst,

Oviposition has been observed in few Aus-

tralian hylids, Watson er al, (1971) described

part of the behaviour associated with egg-lay-

ing in Litorict paraewingt, and | have observed

oviposition in L, citropa, L. wentate, L. frey.

cineti and L. glaserti. Some of the ovipositional

patterns in L, verreauxi ure unique, notably

the action of the male pushing the cluich down

to the fect of the female where the eggs are

held motionless fora short period,

The behaviour of the male in cupping his

feet around the batch and rapidly “fanning” the

eggs may serve to distobute the seminal fluid

around the eggs within a more confined space

and thus aid fertilisation. A similar although

somewhat bnefer process occtirs in the ovi-

positional behaviour of L. citrope, L, déntata

and L, glaverti (Anstis unpubl,), BY holding

the harch still for a period of some seconds,

the female may also aid fertilisation in allowing

time for sperm penetration before the eggs

ure attached to supporting material,

[Lis pat known whether the abdominal con-

tractions in the female prior to egg-laying were

the sole factor in exiruding the eggs, or

whether the pressure exerted by the clasp of

the mije aided the process.

The attachment of eggs to vegetation in

a spiral movement has heen recorder! by

Harrison (1922) for L. verreaext (as Hl.

ewingi) and by Watson er al, (1971) for L.

paraewingi, Warrison's statement that the

female moved “right around the stalk at the

moment of laying" is not borne out by the pre-

sent study, but it is possible that Harrison did

not seethe entire vegtaying procedure. Watson

eral, (1971) state that a female of L, paree-

wingi observed in the field “held onto a sub-

merged grass stem, and pressed the cloaca to

the slem as the eegs were extruded; then the

pair pivoted around the stem while allaching

the eggs”. Such behaviour would appear tobe

similar to that of L, verreauxé except that in the

latter, the female holds the eggs still before

BREEDING BIOLOGY OF LITORIA VERREAUXI aut

alfachment and was not observed pressing the

cloaca 10 the stem during egg extrusion. In the

three oviposition sequences observed im this

species. the extent to which batches of eggs

Were spread around the supporting vevetation

varied, Observations have indicated that the

mortality rate of embryos t Jower in smaller

well-spread batches attached to a stem. Larger

masses of eggs on the bojtam of aquaria with-

out vegetation suffer high mortality from about

stage ¥Y onwards, possibly dwe to inadequate

oxygenation resulting from the thickness of the

ege mass and the depth of the water where

they lay. The attachment of two or three

batches together as one also tends tu increase

mortality, After death of an embryo, a fungus

develops over the egy capsule.

The manner of termination of amplexus

vanes amonust hylids, but often the last ovi-

positional sequence is longer than any other

and is followed by separation cither imme-

diately or a few seconds later, e.g. in Ayla

versicolor (Fouquette & Littlejohn 1960), and

Litoria dentara, £, glanerti and LL. citrapa

fAnstis unpubl.). L. verreauxi also follows this

pattern: however, the brief period of Lota? im-

mobility of both mule and female after separa-

tion has not been recorded in other species.

Ova: The significance of eggs being deposited

in small batches has been discussed by Pyburn

(1963) and Martin & Littlejohn = (1966)-

Harrison's (1922) observation that the eggs

are “attached in a cylindrical mass numbering

upwards of a hundred eggs to grass stalks und

similar submerged objects" 1s probably based

on cases where two or three hatches were

itlached as one,

The ovidiameter in stages 1-8 (1.21 mm)

is in agreement with Harrison’s figure of 1,2

mm. The ovidiarneter of £. ewingi has been

recorded as 1.65 mm (Martin & Littlejohn

1966) and that of L. ewingi and L. veereanst

as 1.7 mm (Martin, Littlejohn & Rawlinson

1966). A scrics of eges of L. ewényt laid in

Adclawte dwnng September 1972, have mean

diameters of 1.18 mm tat stage 1), 120 mm

(stage 5) and |.68 (stages 12-13); measure-

ments similar to embryos of L. verréanxi at the

Same stages (Table 3). I would seem likely

therefore that measurements by Martin ef af.

nay have been taken from embryos at about

Stages 11-13,

The tgesof 4,, paraewitgi are similar to those

of L. ewing) (Watson ef al. 1971). Those of L.

ferviriensis can readily be distinguished from

other members of the complex by the larger

ovidiameter (2.33 at stage 10: Martin & Little-

john 1966). Eges of £. burrowsi can be dis

tinguished from those of the L, ewingt group

by the presence of two jelly layers. around the

ovum. The ovidiameter of this species at stage

14 is close to that of L. jervisienss at the same

stage.

Ermbryes and larvae: The larvae of the L.

ewing? complex are of the common hylid type

(Martin 1967b) as is £, ferrewyi, The draw-

ings by Martin (1967a) of L. burrows! larvae

show u tail not as finely pointed and fins not

as Gdecp as in members of the £. ewirngi com-

plex, The body shape also appears somewhat

different. £. paraewiige larvae are similar to

those of L. ewingi “except ihat the tail fins

(especially the dorsal fin) . . . aré more heavily

pigmented” {Watson e¢ a/, 1971), Specimens

of this species examined are more uniformly

pigmented than L. verreauxi, and three speci-

mens ut stage 26 (mean total length 12.9 min;

body length 7,02 mm) are much smaller than

LL. verreauxi at the same stage. Such size differ-

ences may he relaicd to environmental factors.

The mouthparts of the group are basically

similar, having a formuls of

All have a median gap in the papillae on the

upper lip, the extent of which vartes amongst

individuals of the same species. The number

and size of the papillae is variahle between

species, those of LF. jervisieryis are more

Numerous and tightly grouped than in &,

verreaux?, while those of 1. paraewingi are a

little larger and Jess numerous, The larvae of L.

jervistensis possess larger, darker and more

massive beaks than 2. verreanx? and in a num-

ber of speeriens of the Former species From

Darke’s Forest, the central edge of the upper

heak curves slighly below the level of the rest

of the edge, unlike L, verreauxi. The two pig-

mented areas below the lower beak in L.

verreavixl from Spring Creck, are not found in

other members of the £, ewingi complex.

Larval behaviour ard adepration: All the larvae

of the f£.. ewig? group ate nektonic and

generally exhibit behaviour patterns similar to

those desctibed for 4. verreauxi, However, dif-

ferences occur in the larvae of L. jervisiensis

which have been observed schooling together

in groups of 20 or more in the mid-level of the

water, Individuals fram the group move at

different times (o the surface where they may

take air fAnstis, unpubl.). Larvae of L-

verveaux? were Iever observed congregating in

this manner.

Larval life span and metamorphosis: Data on

larval life span are mainly limited to specimens

in eaptive conditions. Moore (1961) records a

laboratory life spam of three months for EL.

verréaux’ Which agrees with one of the groups

fuised at Penshwrst. Harrison (1922) found

that larvae in aqnaria “required upwards of

three months” lo reach metamorphosis, but

believed seven to eight weeks to be normal life

span in ibe field during summer, This ts con-

siderably less than the approximate minimum

of 79 davs for one group in the present study,

but this difference may simply reflect different

culture and temperature conditions, Further

observations are fecessuty lo ascertain the

average life span of this species in the field,

Moore (1961) records the body lengths of

1! newly metamorphosed L. verreauad as 14.3-

17.00 mm: consistent with measurements of

specimens in the present study (Table 3).

Martin (1965) gives a range 11,J-13.6 mm

MARION ANSTIS

for newly metamorphosed L. swinger, which are

generally smaller than L, verreauxi, and Martin

& Littlejohn (1966) 15.6-19.7 mm for L. jervis-

iensis, No data on B, furrewsi and L. parae-

Wing: are available.

The overall life cycle of L. verreanal appears

quite similar ta that of other members of (he

£. ewingi complex in the adaptations to still

water situations, although L. jervisiensiy differs

noliceably in the details of its life history

(Martin & Littlejahn 1966). More data are

necessary before useful comparisons can be

made between the life histories of £. burrowsi

and the £. ewing! complex.

Acknowledgments

I gratefully acknowledge the receipt of a

Tesearch grant from the C.S.J-R.O. Science &

Industry Endowment Fund, which has greatly

facilitated this study. De Harold Cogver, Mr

Harry Ehmann, Dr Norman Gradwell, Dr

Angus Martin and Me Michael Tyler tread and

constructively criticised the manuseript. Thanks

are also due to Mr and Mrs D. Anstis for

assistance im various ways.

References

Ansris, M. {1976),—Courtship bebaviour in the

Australian tree frog Litoria ewinei ¢Anura:

Hylidae). Merpetofauna (1). 16,

Carianin S, J. (1957),—Australian teee frogs of

tbe genus Ayla. Proe. Linn, Soc. NSW. 82,

~108,

Fuercien, J, F. (1889)—Observations on the

oviposition and habits of certain Australian

Batrachians. Prac: Linn. Soc. N.S.W’, 4, 357-

387,

Fougurtre, M. §., Jr & Littntyonn, M, J-

(1960),—Patterns of oviposition in two

speed of hylid frops. SW. Naturalist 5(2),

-96.

Gosner, K, L. (1960)—A simplified table for

staging anuran embryos and Jarvae with notes

on identification. Herpetofagiva, 16, 183-190,

GRADWELL, N. (1972) —Gill irrigation in Rana

cdteshelana. Pt. 1, On the anatomical basis.

Cun. £. Zoael, 50, 481-499,

Harerson, T,, (1922).—On the breeding habits of

some Australian frogs, Aust. Zoal. 3, 17-34.

Citriesoun, M, J. (1963).—Frogs of the Mel-

bourne area. Mier. Nat. 79, 296-304,

LITTLEJOHN, M. J. (1965).—Premuating isolation

in the WWyla ewingl complex (Anurca:

Hylidae). Evolution 19, 234-243.

Manrin, A. A, (1965)—Tadpoles of the Mel-

bourne area, Tier. Nef. 82, 139-149.

MarTIN, A. A. (1967a).—The exrly development.

of Tasmania’s endemic Anura, with comments

on their relationships. Prec. Linn. Soc. N.S.W.

92, LO7-116,

Marnn, A. A, (1967b}.—Australian anuran life

histories: some evolutionary and ecological

aspects. In A, H. Weatherley (Ed.), “Austra-

lian Inland Waters and their Fauna’, (Aust.

Nat. Univ. Press; Canberra, )

Maatin, AW A. & Lirt.esonn;, MJ, (1966).—

The breeding biology and Jarval development

of Hyla jervisiensixs (Anura: Hylidae), Proc,

Linn, Sac. N.8.H'. 91, 45-57,

Maatin, A. A., Lirriegoun, M. J., & Raw iinsen,

P. A, (1966).—A key to the anuran eges of

the Melbourne area, and un iddition to the

anuran fauna. Vier. Nat. 83, 312-315

Marin, A. A. & Watson, G, F. (1971),—Life

history as an aid to generic delimitation in

the family Hylidae. Cepeia, 1971(1), 78-89.

Moore, J. A. (1961).—The frogs of eastern New

South Wales, Bull, dm. Mux, Nat, Afst, 124,

149-386.

Orton, G, L, (1953).—The systematics of verte-

brate larvae. Syst. Zool. 2(2), 63-75.

Pysunn, W. FP. (1963}—-Observalions on the life

history of the tecefrog, Ployllomedusa calli-

dryas (Cope). Texas J. Sei. 15, (S5-170,

Tyree, M, J, (19713,—The phylogenetic signiti-

cance of Vocal sac structure in hylid frogs.

Brin Ransus Publ. Max. Nut. Mist. 19, 319-

360.

Watson, G. FB. Lorrus-Hitus, J, J. & brrerur-

gous, M. J, €1971).—The Litera ewinel

complex (Anura: Hylidae) in south-castern

Australin 1. A new species from Victoria.

Awst, J. Zool, 19, 401-416,

RESTRICTION OF THE CHIRIDOTID GENUS TROCHODOTA LUDWIG

(1891) (HOLOTHURIOIDEA: APODIDA), WITH THE DESCRIPTION OF A

NEW SPECIES FROM SOUTH AUSTRALIA

BY F. W. E. ROWE*

Summary

ROWE, F. W. E. ( 1976) .-Restriction of the chiridotid genus Trochodota Ludwig (1891)

(Holothurioidea: Apodida), with the description of a new species from South Australia. Trans. R.

Soc. §. Aust. 100(4), 203-206, 30 November, 1976.

Trochodota Ludwig (1891) is restricted to the type-species T. purpurea (Lesson), and three other

species, including a new species from South Australia. The generic significance of scattered or

heaped wheels, used in separating Trochodota from Taeniogyrus Semper, 1868, is disputed. The

distribution of serrations on the inner margin of the wheels is regarded a more reliable generic

character; on this basis seven species included in Trochodota by H. L. Clark (1921) and subsequent

authors are referred to Taeniogyrus.

RESTRICTION OF THE CHIRIDOTID GENUS TROCHODOTA LUDWIG (1891)

(HOLOTHURIOIDEA: APODIDA), WITH THE DESCRIPTION OF A NEW

SPECIES FROM SOUTH AUSTRALIA

by F. W. E. Rowe*

Summary

Rows. F. W. E. (1976)—Restriction of the chiridotid genus Trocliidota Ludwig (1891)

(Holotharicidea: Apodida), with the description of a new species from South Australia,

Trans. R. Soe, S$, Aust, 100(4), 203-206, 30 November, 1976,

Trochodota Ludwig (1891) is restricted to the type-species J. purpurea (Lesson), and

three other species, including a new species from South Australia, The generic significance of

scattered or heaped wheels, used in separating Trovhodota from Taentogyrns Semper, 1868,

is disputed, The distribuiion of serrations on the inner margin of the wheels is regarded a

more reliable generic character; on this basis Seven species included in Trochodumm by H. Li

Clark (1921) and subsequent authors are referred to Tdenlogyrus,

Introduction

Among the holothurians collected at Port

Lincoln, South Australia during 1975 by Mr

S, A. Shepherd are six belonging to an undes-

embed species congeneric with Trochodota pur-

puree (Lesson), type-species of Trochodota

Ludwig (1891).

In one important character, the new species

falls into an intermediate position between the

genera Trochxlota and Taeniogyrus Semper

(1868), as currently diagnosed (H. L. Clark

1921; Pawson 1964), A review of the two

genera has revealed that they are not based

upon reliable characters; in this paper they are

tedescribed and a list of species given for cach,

It is not appropriate to discuss in this paper

the validity of all species now included in

Taéniogyrus, since maby species require re-

examination and material is not available:

H. L. Clark (1921); Pawson (1964). However,

the differences between the four well docu-

mented species in Trechedota, including the

new species described below, are tabulated.

Taxonomic account

H. L. Clark (1921), revising the chiridotid

genera, separated Tueniogyrus Semper (1868)

and Trochedote Ludwig (1891) from other

genera, because they possess a combination of

wheel and sigmoid ossicles. On the basis of

having the wheel ossicles actually collected into

sharply defined papillae of the body wall,

Taeniogyrus was considered generically distinct

from Trochodota, Small accumulations of

wheels were considered indicative of T'rocho-

dota. Subsequent authors have rigidly adhered

ta this recognition of the two genera (A. M.

Clark 1966; Hickman 1962; Pawson 1964,

1970; Heding (928; Cherbonnier 1952).

Although several new species of Taeniogyrus

have been described since 1921, no new species

of Trechedote have been found,

With the arrangement of wheels in large

groups, though not in papillae, the new species

described below falls into an intermediate posi-

tion between Taeniogyrus and Trechodota. In

my view, this. shows the unreliability of using

such a character for generic distinctions, par-

ticularly when H. L. Clark (1921) used the

similar grouping of sigmoid ossicles for species

duterminations, <A difficulty then arises in

deciding the relative merits of the importance

of wheels versus sigmoid ossicles in the recog-

nition of the generic taxa, for which no sound

argument has so far been advanced. One

eharacter which does so easily distinguish T.

purpurea, T. allani, T. maculata and the new

species nol only from those in Taentopyrus, is

the arrangement of the serrations on the inner

margin of the wheels. In the absence of any

other reliable internal or skeletal character, 1

believe that this is a much more significant

* Australian Museum, P.O. Box A285, Sydney South, N.S.W, 2001.

character on which to place generic weight. ft

also accords with the use of spicule form in the

recognition of generic taxa within other orders

of holothurians (Panning 1949; Rowe 1969),

Tréchedota is herein restricted to that dis-

crete group of species with serrations of the

inner tim of the wheels arranged in groups.

The remaining seven species included by H. L.

Clark (1921) in Trochodota are referred to

Taerniogyras.

Trochodata is now considered to be

restricted to the southern hemisphere, with rep-

resentative species ranging from the colder

waters of the southern tip of South America

to the more temperate waters of southeastern

Australia, including ‘Tasmania, and the tropical

waters of the Torres Strait between Papua New

Guinea apd the northeastern tip of Australia.

The genus is found from shore-line to depths of

about 50 m.

Taeniopyrus Scmper, 1862

Taeniogyrus Semper 1868: 23.

?Sipmodota Studer 1876: 454,

?Trechodota Ludwig 1891: 358 (part).

Oiagnosts: Chiridotid genus with wheels and

sigmoid ossicles present, scattered, or in groups

or clustered into papillae; wheels with serra-

tions continuous around the inser margin: ten-

tasles LO or 12,

Type-species: Chiridota australianus Stimpson

1836,

Other species: T. contortus (Ludwig 1874); T.

cidaridis Oshima 1915; T, dubius H. L. Clark

1921, T_ ketensis Heding 1928; T. clarus

Heding 1928; T. dunedinensis (Parker 1881);

T. diasema (H. L. Clark 1921); T. roebucki

(Joshua 1914); 7. rosea (Oshima 1914); T,

japenica (von Marenzeller 1881); T. dendyi

(Mortensen 1925): T. dayi (Cherbonnier

1952); 27. venusta Semon 1887.

Trochodota Ludwig, 1891

Trochodota Ludwig 1891: 358 (part).

Diagnosis: Chiridotid genus with wheels and

sigmoid ossicles present scattered or in groups,

Wheels with serrations on the inner margin in

well defined groups; tentacles 10.

Type-species: Holothuria (Fixstulariu) purpurea

Lesson 1830.

Ocher species: T. allani (Joshua 1912); T,

maculata A. L. Clark 19215 7. shepherdi n. sp,

F. W. BE, ROWE

Figs 1-4. Travthodoia shepherdi mep. Fig.

Tentacle rods; Tig. 2—Two radia! and

two interradial plates of the calcareous

ring: Fig. 3—Wheel ossicle; Fig, 4—Sig-

moid ossicle,

1—

Trochodota shepherdi n, sp.

T. allani, Joshua & Creed, 1915: 2! (non 7.

allani Joshua).

Types: Holotype (Australian Museum J9467)

and 5 paratypes. J9796 (2) J9797 (1): Sauth

Australian Museum K1366 (2)); Proper Bay,

Port Lincoln, Spencer Gulf, S. Aust., among

algae growing on Pinna dalobrata at 10 m™

depth. Collected by S$. A. Shepherd,

23.vill.1975,

Diagnosis: Large spicules, wheels. 55-226 xm

diameter; serrations on inner margin of whecls

in G groups of about 16; sigmoid ossicles 144—

190 um Jong, outer curve of hook of sigmoid

ossicle with minute thorns; colour in hfe, black,

Description of holatype (Which has been cis-

sected): The holotype is 60 mm long, of which

the anterior 20 mm is contracted. Body dia-

meter 5-8 mm along its length, tapering only

at the posterior end, Ten strongly contracted

tentacles, each with about 3 pairs of digits.

NEW SPECIES OF TROCHODOTA 205

TABLE 1

Differences between species of Trocvhodata

Species Siemoid ossicles Wheel ossicles Colour Distribution

Grouping Hook Length Grouping Diameter

meculate in papillae and smooth 66-77 um scattered 50-100 wm = pink Torres Strait,

scattered with Qld

darker

spots

allani T ‘Scattered per- smooth 120-150 wm scattered singly 33-216 yum purple Port Phillip,

pendicular ta of at most io Vic—-S.E. Tas.

longitudinal small groups of

body axis 5-6

purpurea scattered smooth 125-150 um scattered 184-182 ym purple Southern

Ocean, Falk-

land Istand

southern coast

of South

America

ahepherdi scatiered per- swith 144-190 2m discrete, large 84216 am black Port Lincoln

pendicular to thorns groups (not in and Kangaroa

longitudinal papillae) 1, S. Aust.

body axis arranged uni

serially along

each inter-

radius

Spicules in tentacles comprise slightly curved

rods dichotomously branched at each end, and

usually have a series of thorny knobs projecting

laterally along shaft (Fig. 1).

Calcareous ring comprises 10 pieces fused,

with .@ straight ianterior and a slightly wndulat-

ing posterior margin, Radial pieces each have a

small anterior notch, Each piece of the ring is

1 mmx 0.5 mm (Fig. 2).

There is one ventral polian vesicle and the

dorsally placed madreporite ig very small and

hook-shaped (0.5 mm x 0.25 mm)-

Long. unbranched gonad on either side of

the dorsal mesentery, jojned anteriorly to a

single, dorsal gonoduct. Gonads extend for

about two-thirds of the body length and are

pucked with eggs.

Ciliated funnels numerous: on either side

ut base of dorsal mesentery in mid-dorsal inter-

radius in mid-interradial line of interradius ad-

jacent to, and to left of mid-dorsal intcrradius,

and in ventral interradius directly opposite to

those two dorsal interradii, Body wall trans-

lucent; radial muscles, lines of ciliated funnels,

and outline of internal organs can be seen

through it.

Calcarcous spicules of hody wall comprise

Wheels and sigmoid ossicles. Wheels restricted

to discrete groups but not accumulated into

papillae. These groups form a single line along

each of the interradii, except in the posterior

1/3 of body where the groups possess smaller

numbers of wheels, and form two irregular

lines. per interradius. Wheels have six spokes,

Inner margin of each wheel has six discrete

groups of about 16 serrations (Fig. 3). Wheels

are 35 pm-226 um in diameter. Sigmoid

ossicles evenly scattered throughout the body

wall and lic perpendicular to longitudinal axis

of body. Shaft of each is smooth except that on

the outside curve. at the attenuated hook end,

there are 2-4 minute spines or thorns (Fig, 4).

The sigmoid ossicles are 144-190 4m in length.

The animal im life is black. Besides the holo-

type 5 other specimens, similar in all respects,

Were collecled and these are considered as

paratypes.

Distribution: Port Lincoln, Spencer Gulf and

Kangaroo Island, South Australia.

Etymology: The species is named after the col-

lector, Mr Scoresby A. Shepherd.

Remarks. Differences between T. shepherdi, T-

maculata, T. allani and T. purpurea are listed

in Table 1.

Acknowledgments

I would like to thank Mr S, A, Shepherd

(Department of Fisheries, South Australia) for

forwarding material of the new species to me

206

for examination; Dr 8B. J. Smith, National

Museum of Victoria; and Mr W, Zcidler, South

Australian Museum, for allowing me to exa-

F, W. E, ROWE

mine remaining slides and specimens of T.

allani and 7. roebucki described or examined

by Joshua (1912, 1914),

References

CHERBONNIER, G. (1952),—Contribution a la con-

naisance des Holothuries de l'Afrique du Sud.

Trans. R. Soc. §, Afr. 33(4), 469-509, pls

XXXV-L.

Crark, A. M. (1966).—Echinodermata, Port

Phillip Survey. Mem. natn. Mus. Vict. 27,

289-384, pls i-iv,

Ciark, H. L, (1921).—The cchinoderm fauna of

Torres Strait. Pap. Dep. mar. biol, Carnegie

Instn, Wash. 10, 1-223, pls 1-38.

Hepinc, S. G. (1928)—Synaptidae. Vidensk.

Meddr dansk naturh. Foren. $5, 105-323, 69

figs, pls 2-3.

HickmMan, V. V, (1962).—Tasmanian sea-cucum-

bers (Holothuroidea). Pap. Prec, R. Soc.

Tas. 96, 49-72, figs 1-177, 2 pls.

Josuua, E. C, (1912).—On a new holothurian of

the genus Taeniogyrus found in Port Phillip

Bay. Proc. R. Soc. Vict. (1.8.) 25, 79-81, pls

MAyV.

Josuua, E. C. (1914).—Victorian Holothuroidea

with descriptions of new species. Proc. R. Soc.

Viet. (ns.) 27(1), 1-1], pl. 1.

Josuua, E. C. & Creep, E. (1915).—South Aus-

tralian Holothuroidea, with descriptions of

new species. Trans. R. Soc. §. Aust, 39, 16-24,

3. pls.

Lupwic, H. (1889-1892), — Echinodermen

(Stachelhauter). 1. Buch. Die Seewalzen. Jn

Bronn, H, G. Klassen und Ordnungen des

Thier-Reiches. Leipzig 2(3), 1-447, 25 figs,

17 pls, 12 maps,

Mortensen, T. (1925).—Echinoderms from New

Zealand and the Auckland-Campbell Islands.

Hl Asteroidea, Holothurioidea and

Crinoidea. Vidensk. Medd. naturh. Foren.

Kbh. 79, 263-420, 70 figs, 3 pls.

PANNING, A. (1949).—Versuch einer Neuordnung

der Familie Cucumariidae (Holothuroidea,

Dendrockitota), Zool. Jb,, 18, 404-470, 62

gs.

Pawson, D. L. (1964).—The Holothuroidea col-

lected by the Royal Society Expedition to

Southern Chile, 1958-59. Pac. Sci. 18(4),

453-470, 3 figs.

Pawson, D, L. (1970).—The marine fauna of

New Zealand: Sea cucumbers (Echinoder-

mata: Holothuroidea). Bull. N.Z. Dep,

scient. ind. Rés. 201, 7-69, 10 figs, 2 pls.

Rowe, F, W. E. (1969).—A review of the family

Hofothuriidae (Holothurioidea: Aspidochiro-

tida). Bull. Br. Mus. nat. Hist. (Zool.) 18(4),

117-170, 21 figs,

SEMPER, C. (1868).—Die Holothurien. Reisen im

Archipel der Philippinen. Weisbaden 1, 228

pp., 60 pls.