(Registered at the General Post Office, Perth, for transmission by post as a Periodical.}

JOURNAL

OF

THL ROYAL SOCIETY

OF

WL5TLRN AUSTRALIA, INC.

Founded 1913 :: :: Incorporated 1937

voi. xxvin

1941 - 1942

The Authors of Papers are alone responsible for the statements

and

the opinions expressed therein.

Published 22nd May, 1944,

Printed for the Society by
Robbbt H. MmLER, Government Printer, Perth.

1944.

pr:t/rr-

[ Registered at tlie General Post Office, Perth, for transmission by post as a Periodical.]

JOURNAL

OF

THL ROYAL 50CILTY

OF

WESTERN AU5TRAEIA, INC.

Founded 1913 ;; Incorporated 1937

Vol. XXVIII

1941 - 1942

4’he Authors of Papers are alone responsible for the statements

and

the opinions expressed therein.

Published 22nd May, 1944.

Printed for the Society by
Robkiit H. Miller, Government Printer, Perth.

1944.

k 119/43

11.

Jour. Roy. Soc., Western Australia, Vol. XXVIII.

THE ROYAL SOCIETY OF WESTERN
AUSTRALIA, Ixc.

OFFICERS AND COUNCIL, 1941-42.

Patron :

His Majesty the King.

Vice-Patron :

His Excellency Sir James Mitchell, K.C.M.G.,

Lieut. -Governor of the State of Western Australia.

President :

C. A. Gardner.

Past President :

^ Professor A. D. Ross, M.A., D.Sc., F.R.S.E., F.R.A.S., F.I.P.

Vice-Presidents :

R. C. Wilson, B.Sc., B.E. E. M. Watson, B.Sc., Ph.D., D.I.C., A.I.C., F.A.C.I.

Joint Hon. Secretaries :

R. T. Prider, B.Sc., Ph.B., F.G S. 0. F. H. Jenkins, M.A,

Hon. Treasurer :

L. J. H. Teakle, B.Sc. (Agric.), Ph.D., A.A.C.I.

Hon. Editor :

B. L. Southern, A.A.C.I.

Hon. Librarian ;

Eileen A. Jenkins, M.Sc.

Hon. Asst. Librarian :

Dorothy Carroll, B.A. B.Sc., Ph.D., D.I.C.

Council :

Allison M. Baird, B.Sc.

H. W. Bennetts, D.V.Sc.

H. Bowley, F.A.C.I.

Professor E. de C. Clarke, M.A.

F. G. Forman, B.Sc.

L. Glauert, B.A., F.G.S.

R. J. Little.

H. P. Rowledge, A.A.C.I., A.W.A.S.M.

Hon. Auditors :

J. Shearer, B.A., M.Sc. W. F. Cole, B.Sc.

Jour. Roy. Soc. Western Australia, Vol. XXVIII.

iii.

CONTENTS.

Volume XXVIII.

Page

Office Bearers ... ... ... ... ... ... ... ... ... ... ii.

Annual Report ... ... ... ... ... ... ... ... ... ... iv.

Treasurer’s Report ... ... ... ... ... ... ... ... ... ... vi.

Proceedings — Abstract of ... ... ... ... ... ... ... ... ... vii.

Index of Authors ... ... ... ... ... ... ... ... ... ... x.

General Index Ixxxix.

Presidential Address — The Vegetation of Western Australia by C. A. Gardner ... xi,

1. Permian Productinae and Strophalosiinae of Western Australia. By K. L. Pren-

dergast ... ... ... ... ... ... ... ... ... ... 1

2. The Occurrence of the Genus Conodypus in the North-West Division, Western

Australia. By Irene Crespin ... ... ... ... ... 75

3. Fibroferrite and Copiapite from Yetar Spring near Chidlows. By R. T. Prider 79

4. The Geology and Petrology of part of the Toodyay District, Western Australia.

By R. T. Prider 83

5. Revision of the Embioptera of Western Australia. By Consett Davis ... ... 139

6 . Ecological Succession observed during Regeneration of Triodia pungent, R. Br.

afterburning. By Nancy T. Burbidgs ... ... ... ... ... 149

7. Ecological Notes on. the Vegetation of the 80-mile Beach. By Nancy T. Burbidge 157

8 . Bryozoa from the Wandagee and Nooncanbah Series (Permian of Western

Australia) — Part I. By Joan M. Crockford 165

9. A Revision of some previously described species of Bryozoa from the Upper

Palaeozoic of Western Australia. By Joan M. Crockford ... ... ... 187

10. Fossil Plants from Gingin, Western Australia. By A. B. Walkom 201

11 . Marine Copepods from Western Australia (Series II.). By W. S. Fairbridge ... 209

12. A Consideration of the Insect Population associated with Cow Dung at Crawley,

W.A. By G. J. Snowball ... 219

13. The Essential Oils of the Western Australian Eucalypts. By E. M. Watson ... 247

IV.

Jour. Roy. Soc., Wp:stp:rn Australia, Vol. XXVIII.

The Royal Society of Western Australia (Inc.).

ANNUAL NEPORT OF THE COUNCIL FOR THE YEAR ENDING

30th JUNE, 1942.

Ladies and Gentlemen,

Your Council begs to submit the following report for the year ended
30th June, 1942.

Council . — As in the past several years an Executive Committee was ap-
pointed to deal with the routine business of the Society and to report quar-
terly to the Council. The Council met on six occasions and the Executive
Committee on six occasions during the year.

Finance . — The General Fund shows a. balance of £146/1/5 and the
Endowment Fund now amounte to £271. There are certain commitments to
the Government Printer against the General Fund in connection with the
publication of Volume 27, which has not yet been completed.

Membership . — Membership remains almost the same as at the commence-
ment of the year. Members elected during the year include one Correspond-
ing member, eight ordinary members and two associates, while two associates
have transferred to full membership. Six ordinary members have resigned
during the year.

We regret to record the loss by death of four of our members — Mr. W.
E. Shelton, Drs. W. E. Blackall and H. J. Lotz and the Hon. John Nichol-
son. All of these gentlemen took a keen personal interest in the proceed-
ings of the Society. Mr. Shelton was at various times president, librarian
and secretary of this Society. The Hon. John Nicholson acted for some years
as a member of the Endowment Lands Committee and rendered the Society
considerable assistance at the time of its incorporation.

There are at present 150 members of the Society made up as follows: —

Honorary meml)ers . . . . . . . . 7

Corresponding members . . . . . . 9

Life member . . . . . . . . ■ • 1

Ordinary members . . - - • • • • 98

Associate members • • • • • • • • 35

Journal . — Volume 27 is nearing completion, and a number of the papers
for the succeeding volume, No, 28, are in course of publication. During the
past year fourteen ))apers were ]iresented to the Society for publication in
the Journal.

Jour. Roy. Soc., Western Australia, Vol. XXVIII.

V.

Mr. Southern is relinquishing the ofhce of Editor after twelve years'
service in this capacity. The Council wishes to record its appreciation of
Mr. Southern's excellent work in connection with the publication of the
Journal during this period. Mrs. Jenkins is undertaking the duties of
editor.

Library.— The Society has now entered into exchange of publications
with a total of 189 institutions, of which 56 are in Australia, 16 in the United
Kingdom, 23 in other parts of the British Empire, 48 in North and South
America, 42 in Europe and four in Asia. Many exchange publications are
coming to hand fairly regularly, but in some instances overseas institutions
are holding their publications in reserve to be forwarded after the war.
The Council of this Society has decided, in view of the danger of loss during
transit, to hold copies of the Journal which would normally be forwarded
to overseas exchanges and to forward them in more peaceful times.

The Council wishes to express its thanks to Mr. A. Gibb Maitland for
further donations to the Society's libraiw made during the year.

C. A. GARDNER,

President.

REX. T. PRIDER,

C. E. H. JENKINS,

Joint Hon. Secretaries.

THE ROYAL SOCIETY OF WESTERN AUSTRALIA, INCORPORATED.
Statement of Receipts and Expenditure for the Year ended ‘ 6 t)th June. 1042.

vi. Jour. Roy. Soc., Western Australia, Vol. XXVIII.

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Amounts Owing — Rent .... .... .... 7 10 0 General Fund .... .... .... .... 146

Investment — Life Membership Fund .... .... .... .... 18 0 0 Medal Fund .... .... .... .... .... .... .... 4

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viii. Jour. Roy. Soc., Western Australia, Vol. XXVIII.

ABSTRACT OF PEOCEEDINOS, 1941-1942

8th July, 1941 —

Annual General Meeting in Gleclclen Hall. Presidential Address: “Develop-
ments in Optical Science/’ by Professor A. D. Ross, M.A., D.Sc.

Royal Society’s Medal — Presentation of the Royal Society’s Medal by His
Excellency the Lieutenant Governor, Sir James Mitchell, K.C.M.G., to
Professor E. de C. Clarke, M.A.

I 2th August, 1941 —

Po/per — “The Petrology of Part of the Toodyay District,” by Dr. R. T.
Prider.

Talk — ■“Detection of Forest Fires,” by Mr, A. C. Shedly, M.Sc.

9th September, 1941 —

Paper — “The Occurrence of the Genus Conoclypus in the North-West Divi-
sion, Western Australia,” by Irene Crespin, B.A., communicated by
Professor E. de C. Clarke, M.A.

“Insects and War,” by Mr. C. F. H. Jenkins, M.A.

Exhibits — Dr. R. T. Prider exhibited specimens of gypsum crystals collected
from the walls and roofs of old workings at Oroya North Blocks and
Paringa Mines, Kalgoorlie.

14th October, 1941 —

Paper — “ Fibroferrite and Copiapite from Yetar Spring,” by Dr. R. T.
Prider.

Talk — ”How the Flour Mill Works,’’ by Dr. L. W, Samuel.
llTPi November, 1941 —

Talk — “Copper Deficiency Diseases in animals in the South-West,” by Dr.
H. W. Bennetts.

Exhibits — Mr. L. Glauert exhibited portions of a skull of the extinct mar-
supial, JHprotodon. The specimen -was collected by Mr. L. L. Miller,
of Warrawagine Station, Oakover River.

9th December, 1941 —

Papers — “Regeneration of Triodia pnngens, R.Br. after burning with special
reference to ecological succession,” by Nancy Burbidge, B.Sc. (Hon.).

“Revision of the Embioptera of Western Australia,” by Mr. Consett Davis,
M.Sc., communicated by Mr. L. Glauert, B.A.

10th IVIarch, 1942 —

Papers — “Ecological Notes on the De Grey-Coongan Area with special refer-
ence to Physiography,” by Nancy Burbidge, B.Sc. (Hon.).

“Ecological Notes on the Vegetation of the 80-mile beach,” by Nancy Bur-
bidge, B.Sc. (Hon.).

Talk — “W.A. Stone Implements,” by Mr. H. V. V. Noone, M.R.A.I.,
M.R.A.S.

Exhibits — Mr. A. Gibb Maitland exhibited some fine examples of New
Guinea stone implements and drew attention to the different type of
workmanship compared with the exhibits illustrating Mr. Noone ’s
address.

•loUK. Roy. 8oc., Western Australia, Vol. XXVIII.

\x.

14tii April^ 1042—

Papers — ‘ ■ Bryozoa from tlio WamlaoTO aiul Nooiicanbah series of Western
Australia,” Part I, by *loan Croekford, M.Bp., eomrnimieated by Pro-
fessor E. de 0. Clarke, M.A.

“A Revision of some previously described species of Bryozoa from the
rapper I'alaeozoic of Western Australia,” by Joan Croekford, M.Sc.,
communicated by Professor E. de C. ('larke, M.A.

“Fossil Plants from Oingin, Western Australia,” by Dr. A. B. Walkom,
communicated by Professor E. de C. Clarke, M.A.

‘‘Two New Co])epods from Western Australia,” bv Mr. W, B. Pairbridge,
B.Rc.

12t!i AIay, 1942—

Papers — “Metamorphosis of tlie Jas])er Bars of Western Australia,” by
Or. K. R. Miles.

E.rhihits — Dr. R. T. Prider exhibited aji example of Banded Quartz Hypers-
thene Rock from 12 miles north-west of Bolgart.

Oth Jt'NE, 1042 —

Papers — “A consideration of the insect Population associated with Cow
Dung at Crawley, W.A.,” by Air. C. J. Snowball, B.Bc. (Ifon.).

“The Essential Oil of Eucalppins erythremewa^’ by Dr. E. M. Watson.

Exhibits — Mr. R. C. Wilson exhibited specimens of copper from Marda as
an exani})le of ihe good results obtained by the use of small smelting
plants.

X.

Jour. Boy. Soo., Western Australia, Vol. XXVTIT.

INDEX OP

Burbiflge, N.

Crespiii, T.

Davis, C

Faivbvidge, W.

Gardner, C. A.

Prendorgast, K. L. ' . .

Prider,, P. T '

Croekford, .T.

Snowball, G. J.

Walkom, A. B.

Watson, K, M.

AUTHORS.

149 , 157

75

139

209

xi

1

! . . . 79 , 83

165 , 187

219

201

247

The Vegetation of Western Australia.

XI.

].— THE VEGETATION OF WESTERN AUSTRALIA.

WITH SPECIAL REFERENCE TO THE CLIMATE AND SOILS.

PRESIDENTIAL ADDRESS, 1942.

By Charles Austin Gardner.

Read 13th July, 1942 ; published 28th March, 1944.

CONTENTS.

Page

I. Introduction .. . .... .... .... .... .... .... .... xii.

II. The Elements of the Australian Flora .... .... .... .... xiii.

1. The Antarctic Element .... .... .... .... .... xv.

2. The Palaeotropic Element .... .... .... .... .... xx.

3. The Australian Element .... .... .... .... .... xxiii.

III. Physiography ... .... .... .... .... .... .... xxvii.

IV. Geology and Soils .... .... .... .... .... .... .... xxix.

V. Climate .... .... .... .... .... .... .... .... xxx.

1. The Climatic Cycle .... .... .... .... .... .... xxx.

2. Temperature .... .... .... . .. .... .... .... xxxiii.

3. Humidity, Evaporation and Wind .... .... .... xxxiv.

4. Ratio of Precipitation to Evaporation .... .... .... xxxiv.

VI. Relationships between the Climate and Vegetation of Western

Australia .... .... .... .... .... .... .... .... xxxv.

VII. The Provinces and their Formations .... .... .... .... xxxviii.

1. The Northern Province .... .... .... .... .... xxxviii.

2. The South-West Province .... .... .... .... .... xli.

(a) The Forest Formations .... .... .... .... xlii.

(i) The Jarrah Forest .... .... .... .... xlii.

(ii) The Karri Forest .. . .... .... .... xliii.

(iii) The Tuart Forest .... .... .... .... xliii.

(b) The Woodland Formations .... .... .... .... xliv.

(i) Temperate Savannah Woodland .... .... xliv.

(ii) The Sclerophyllous Woodland .... .... xliv.

(c) Lateritic Formations .... .... .... .... xlvi.

(d) Formations of the Granite Rocks .... .... .... xlvi.

(e) Swamp Formations .... .... .... .... ... xlvii.

(f) Halophytic and Dune Formations .... .... .... xlviii.

(g) Sand-heaths .... .... .... .... .... .... xlviii.

3. The Eremean Province .... .. . .... 1.

The Formations .... - .... .... .... .... .... li.

(a) Sclerophyllous Woodland .... ... .... .... li.

(b) Salsolaceous Shrub Steppe .... .... .... li.

(c) Mulga Bush .... .... .... .... .... li.

(d) Halophytic Formations .... .... .... .... liv.

(e) The Triodia Steppe .... .... .. . .... Iv.

(f) The Desert .... . .. .... . .. .... .... Ivi.

Xll.

Charles Austin Gardner.

VIII, Epharmosis and Growth-forms

1. Trees and Shrubs

2. Lianes ....

3. Herbs ....

Ivii.

Iviii.

Ixi.

Ixii.

IX. Climatic Tables —

Table I. — Mean Monthly and Annual Precipitation in Points

(100 points ^ one inch) .... .... .... .... Ixvi.

Table II. — Annual and Seasonal Rainfall Tables, showing
Rainfall for the four consecutive wettest months, the four
consecutive driest months, the percentage of the four
wettest months, and the Annual Rainfall Variation ex-

pressed as a percentage of the annual rainfall .... .... Ixix.

Table III. — Relative Humidity — Mean Monthly Readings for
9’0 a.m. and 3*0 p.m. .... .... .... .... . .. Ixxii.

Table IV. — Saturation Deficit — Mean Monthly Averages for

9*0 a.m. Ixxiii.

Table V. — -Climatic Statistics of Rainfall and Temperature,

Saturation Deficit, etc. .... .. . .... . .. .... Ixxv.

X. Maps — ■

Plate I. — Summer Rainfall — November to April — Isohyets

in inches) .... .... .... .... .... Ixxvi.

Plate II. — ^Winter Rainfall — May to October — -(Isohyets

in inches) .... .... .... .... .... Ixxvii.

Plate III. — ^Normal Mean Temperature — Isotherms in ° F.

for January .... .... .... .... .... Ixxviii.

Plate IV. — Normal Mean Temperature — Isotherms in ° F.

for July .... .... .... .... .... Ixxix.

Plate V. — Rainfall Variabihty — Variation from the Mean,

expressed as per cent. .... .... .... Ixxx.

Plate VI. — The Meyer Ratio .... .... .... .... Ixxxi.

Seasonal Precipitation ^

Plate VII.— The " Ratio . Ixxxii.

Seasonal Saturation Deficit

_ _ Seasonal Precipitation ^

Plate VIII. — The Ratio .... .. . Ixxxm.

Seasonal Temperature

Plate IX. — The Three Provinces of Vegetation and Climate Ixxxv.

Plate X. — The Phytogeographical Formations .... .... Ixxxvii.

I. INTRODUCTION.

The geographical distribution of plants and the study of plant habitats
is one of the most interesting aspects of modern botany, and of the countries
of the world, few are more favourably situated for this study and its bearing
on questions of evolutionary history than Western Australia. In few countries
has the ecological aspect been of such general application for, in the general
mosaic of its soil types, the use of plants as indicators of the soil has been
recognised from the eai’liest days of our history as criteria in the selection
of land for agricultural purposes.

The Vegetation of Western Aostralia.

xia.

In a young country such as Western Australia, there are many oppor-
tunities for the botanist and, although botanical investigation of necessity
rests on the foundation of systematic work conducted in the arid environment
of the herbarium, there are many vital problems which can be solved only in
the more inspiring atmosphere of the field. Given this necessary systematic
basis, many are the avenues of approach which are opened up for scientific
research which, because of the almost virgin condition of so much of the
terrain, of^er possibilities denied to workers in the less fortunate older lands.

Phytogeographical research in Western Australia may be said to have
commenced with the publication of Diels’ Die Pflanzenwelt von West- Austr alien
(1906). The foundations, however, were laid down by Sir Joseph Dalton
Hooker in 1859, by the publication of his classic Introduction to the Flora of Tas-
7nania, in which he propounded certain theories regarding the origin and
affinities of the Australian flora, and gave floristic analyses which are invalu-
able for students today. Diels, who visited Western Australia in 1901 and
1902, dealt only with the vegetation of temperate Western Australia. It
must be remembered, however, that at this time, means of transport were
not what they are today, and it is surprising that he accomplished so much
in so short a space of time. The work is, however, an indispensible one for
all students of the plant geography of South-Western Australia.

For over a century, in fact from the time when Dampier visited these
shores, the flora of Australia has claimed the attention of botanists through-
out the world. This is due to the fact that this flora, as a separate geographi-
cal entity, ranks high amongst those floras which are remarkable for the high
degree of endemism amongst their species. Only in South Africa, especially
in the Cape Peninsula, and in South America, do we find anything comparable
in the number of autochthonous species so rich in vegetative and floristic
forms. The Cape Peninsula and South-Western Australia afford interesting
examples for comparison.

It is not that the families comprising the Australian flora are in them-
selves endemic. Far otherwise : out of the total number of these there are
but three families which are strictly endemic in Australia, with a total number
of less than 30 species. Rather is it the number of endemic genera and species,
which is surprisingly large in some areas (especially in the South-West of
Australia), making up as much as 75 per cent, of the total species. There
are a number of families and groups, on the other hand, which are mainly
Australian, that is, the greater number of genera and species are endemic
here, but some of the genera and species also occur in other lands, or, there
are conjunctive genera and species present in these remote areas. These
furnish important evidence, for they stand today as indisputable proof that
tliese widely dispersed areas had at one time a close relationship, if not a
common origin. The presence of these groups and species in countries like
America and Africa, explain two important facts : (a) that the Australian

continent was not always isolated, and that (b) the isolation has been of suf-
ficient duration to enable the distinctive elements of the Australian flora to
make a peaceful and progressive evolution free from any external influences,
resulting in a specialised flora with a degree of endemism rare in other con-
tinents.

II. THE ELEMENTS OF THE AUSTRALIAN FLORA.

The geographical isolation of the Australian continent, its uniform topo-
graphy, and the resultant gradation of climatic areas which range from tropi-
cal to temperate, are the main considerations which must be borne in mind
when we approach the study of the vegetation of Western Australia. Separated

XIV.

Charles Austin Gardner.

from the rest of the world by the wide expanses of the Pacific, Indian, and
Southern oceans, the only geographical approach for terrestrial plants is by
way of Malaya and New Guinea, a line of communication which is interrupted
by the barrier of the Timor and Arafura seas, and the still more restricted
barrier of the Torres Strait. Australia is, therefore, of all the great land
masses of the earth, the most isolated, and it is this isolation that accounts
for its peculiar flora and fauna.

That this isolation did not always exist is evidenced by certain relation-
ships with the vegetation of lands now far removed, such as South America,
South Africa, Malaya, and Madagascar. These relationships are exhibited
by those families and groups which are more or less common to two or more
of these areas. There are certain large families, such as the Compositae, Grami-
neae, and Cyperaceae, which have a more or less cosmopolitan distribution but,
on the other hand, it is amongst the more restricted groups, most highly
developed in Australia — but with relationships with other countries through
conjunctive genera — that these relationships are most clearly exhibited.
To give a general indication of these relationships we may take, by way of
example, the ancient group Gymnospermae. Here we And Araucaria linking
South America w'ith Australia by way of New Zealand and New Caledonia ;
CalUtris and Actinostrohus of Australia have their counterpart in the African
genus Widdringtonia ; while in Cycas there is a link with Asia, and a secondary
link through Macrozarnia with the African Encephalartos. But when we
consider such families as Proteaceae, Epacridaceae, Restionaceae, Goodenia-
ceae, and Stylidiaceae, we find these relationships even more closely expressed,
both through the general limits of areas of distribution, and through con-
junctive genera, and even species.

While today, in the present stage of our knowledge, it is almost impossible
to attempt to explain the origin of the Australian flora, a floristic analysis of
this flora shows us that it has much in common with the floras of the lands
referred to, and thus we are forced to suppose that there existed some former
land communications between Australia, South America, and South Africa.
Whether the groups which today are predominantly Australian and have
spread through the channels of communication suggested, or whether they
are migrants which have become more abundant and diversified here than
in their original home, is a question that may some day be answered by palaeo-
botanists, but not by present day development.

AVhatever may be said of the origin of the Australian flora, we must
recognise the fact that certain elements have been instrumental in its evolu-
tion. These elements appear to have been recruited from two or three princi-
pal sources : an ancient Antarctic or Subantarctic Element, derived from

a southern source, and exhibited by the existing relationships between the
floras of Australia, New Zealand, Southern (principally Andean) South America,
and South Africa ; and a more recent (?) Palaeotropic Element which has
migrated to Australia by way of India, Malaya, and Melanesia.

Diels ]30stulates a third element— the Australian Element, but an analysis
of this clearly indicates that this so-called Australian Element is in reality
a secondary element at the most, for it appears to have too much in common
with both the Antarctic and Palaeotropic elements to be considered in itself
a true element. In his definition of the Australian Element, Diels states :
“ The Australian Element comprises numerically the majority of the plant
species occurring in Australia. Its groups and genera occur either only m
Australia, and have no near relations outside the continent, or they possess

Thk Vegetatiox op Westerx Australia.

XV.

a few wandering representatives which are evidently very closely connected
with the main Australian stock.” He then enumerates those groups which
contain, inter alia^ the following families : —

Centro lepidaceae

Philydraceae

Proteaceae

Pittosporaceae

Stackhousiaceae

Epacridaceae

Goodeniaceae

Restionaceae

Santalaceae

Phytolaccaceae

Tremandraceae

Halorrhagaceae

Myoporaceae

Stylidiaceae

He omits Byblidaceae and Cephalotaccae, which at the time of his work
were not recognised as distinct families, but apart from these, the only en-
demic family is the Tremandraceae.

In addition, he enumerates several groups and certain genera, amongst
which are : Haemodoraceae-Conostylideae ; Leguminosae-Podalyrieae-

Genistcae ; Acacia ; Rutaceae-Boronieae ; Eupliorbiaceae-^S/eno/ofeeae ;
Dodonaea ; Rhamnaceae-Rhamneae ; Sterculiaceae-Buttnerieae-Laszope^aZae ;
Myrtaceae-Leptospermeae-CAomaeZaiic^^eae. The names in italics (mine) are
those of groups which are strictly endemic. h]xamples of some of these will
be discussed in greater detail.

1. THE ANTARCTIC ELEMENT.

Both Hooker and Diels claimed for the Antarctic Element those plants
which were common to South-Eastern Australia, New Zealand, Tierra del
Euego, and Andean Patagonia. This is indeed a very restricted area of dis-
tribution, and the number of conjunctive genera is very small. Typical of
the element in this restricted sense is Nothofayus. Taking this limited view of
the Antarctic Element, there are, beyond a few genera such as Cenirolepis,
Trichocline^ and one or two others, no representatives of the Antarctic Element

estern Australia. The Antarctic Element lias received a wider treatment
from both Field Marshall 8muts and Sir Arthur Hill, who claim for the ele-
ment ■' those plants which have originated from the ancient lands of the South-
ern Hemisphere, as against those which have migrated from the north.”
Under this definition we can recognise certain larger groups which are confined
generally to the southern lands, such as temperate Australia, Now Zealand,
South America, and South Africa, and this broader conception of the Antarctic
Element reduces considerably the importance of the Australian Element in
the sense employed by Diels. Therefore, leaving aside the more restricted
representatives of the Antarctic Element in its narrower sense, we have to
recognise a number of families and genera wliich have a common southern
distribution : an element either restricted to the southern lands, or most

highly developed there, with occasional representatives which have undergone
a northern migration under conditions which favour their spread. Such are
the migrations of Protea into Abyssinia through the highlands, and of Baeckea
and Haemodoriim into eastern Asia, or of Dracophyllum. into northern Queens-
land. C'onversely, we may consider the soutliern migration of Hhododendrcn
in high altitudes into the Bellenden-Kerr mountains in Queensland.

While it is obviously impossible to dc^al comprehensively with all the
groups which comjjrise the Antarctic Element, a (‘onsideration of some of the
more important will illustrate the general distribution of tliose which are
included in it. In the following tables tlu' numbers in brackets refer to the

XVI.

Charlks Austin Gardner.

number of species ; names indicated by asterisks are those of genera not
represented in Western Australia : —

(a) Restionaceae.

A narthria

(0) S.W. Australia

Ecdeiocolea

(2) S.W. Australia

Hopkinsia

(1) S.W. Australia

Lyginia

(2) S.W. Australia

Phyllocomus*-

(1) South Africa

Loxocarya

(8) S.W. Australia

Onychosepalum

(1) S.W. Australia

Chondropetahmi *

South Africa

Lepyrodia

(7) Southern Austr. and N. Zeal.

A skidosperm a *

(1) South Africa

Dielsia

(1) S.W. Australia

Restio

(110) Mainly S. Africa, a few temp. Austr.

Leptocarpus

(12) temp. Austr., Cochin China, and Chile

Elegia^

(20) S. Africa

Lepidobolus

(3) Southern Austr.

Calopsis*

(10) S. Africa

Canncmiois*

(8) S. Africa

Lamprocaulos*

(2) S. Africa

Chaetanthus

(1) S.W. Austr.

Tharmiochortns*

(25) S. Africa

Staberoha'^

(6) S. Africa

Hypolaena

(2) Southern Austr.

Calorophus

(3) Southern Austr. and N. Zeal.

Antkochortus*

South Africa.

Mastersiella*

(10) South Africa

Loxocarya

(8) S.W. Austr.

Harperia

(1) S.W. Austr.

Hypodisciis*

(15) South Africa

Wildenowia*

(15) South Africa i

Swnmary.

13 genera endemic

in South Africa.

10 genera endemic

in South-Western Australia.

2 genera common

to temperate Australia.

2 genera common

to temperate Australia and New Zealand.

1 genus common

to South Africa and temperate Australia.

1 genus common to Chile, Southern Australia, and Cochin China.

(b) Centrolepidaceae.

Gaiinardia*

(3) temp. S. America, N. Zeal., and Austr.

Alepyrum*

(2) temp. S. America, N. Zeal., and Austr.

Juncella

(3) temp. Australia and N. Zealand

Aphelia

(1) temp. Australia and Tasmania

Brizula

(5) temp. Australia and Tasmania

Centrolepis

(20) temp. Australia, N. Zeal., and South

America

The Vegetation of Western Australia.

xvii.

(c) Epacridaceae.

Sect. I. — Prionoteae —

Prionotes^

(1)

Tasmania

Lehetanthus^

(1)

Patagonia

Sect. II. — Epacrideae-

Sprengelia*

(3)

S.E. Australia

Andersonia

(20)

S.W. Australia

Itichea^

(8)

Tasmania and Victoria

Dracophyllum*

(11)

Eastern Australia, N. Caledonia, and N.
Zeal.

Sphenotoma

(6)

S.W. Australia

Cosmelia

(1)

S.W. Australia

Lysinema

(5)

S.W. Australia

Woolsia*

(1)

Eastern Australia

Epacris*

(30)

S.E. Australia, Tasmania, N. Caled., and
New Zealand

Archeria*

(5)

Tasmania and N. Zealand

Sec. III. — Styphelieae-

—

Styphelia

(12)

temp. Australia

Leucopogon

(140)

temp. Australia (mainly S.W.), N, Zeal-
and, Malaya, and Pacific Islands

Lissanthe*

(3)

S.E. Australia

Cyathodes*

(15)

Tasmania, Victoria, N. Zeal., and Hawai-
ian Islands

Cyathopsis*

(1)

New Caledonia

Monotoca

(7)

tern}). Australia

Acrotriche

(8)

temp. Australia

Coleanthera

(3)

S.W. Australia

Brachyloma

(7)

temp. Australia

Conostephium

(5)

S.W. Australia

Pentachondra*

(4)

mountains of Tasmania and Victoria

Trochocarpa

(6)

temp. Australia

Needhamia

(1)

S.W. Australia

Oligarrhena

(1)

S.W. Australia

Summary (26 genera).

8 South-Western Australia.

5 temperate Australia.

4 South-Eastern Australia.

4 Australia, N. Caled., N. Zeal.

2 Australia, extending to the Hawaiian Islands.
1 endemic in Tasmania
1 endemic in Patagonia.

1 endemic in New Caledonia.

ClIARr.KS ArSTINT CJakdxkr.

xviii.

(fl) Stylidiaceae.

Subfam. Doriatioideao —

Donatia* (2) Antarctic 8. America, New Zeal., and

Tasmania

Siibfam. Stylidioideac —

(4) 3 New Zt'al., I Antarctic 8. America
(4) 3 New Zeal., 1 Tasmania
{ I ) New Zealand

(b) ] temp. Australia, 5 S.W. Australia
(105) 90 in W. Austr. (52 endemic). 12 in North-
ern Australia (9 in W. Austr.), 5 common
to temp(‘rate Australia, 5 endemic in 8.E.
Australia, 3 endemic in Indo-Malaya.

Summary (6 genera).

2 common to New Zeal, and Ant. 8. America.

1 extending to Tasmania.

I common to New Zealand and Tasmania.

1 enflemic in Australia.

1 common to Australia and Indo-Malaya.

PhyllacJmp*
Forsfera*
Oreostylidimn *
Leveyihookia
Stylidhim

Subfam. Persoonioid(‘ae
Symph yonem a *
Belle7idena*
Agastachya*

Beauprea *

Dilobeia'^

Cenarrhenes"^

Garnieria*

Persoojiia

Brabeiuni*

Subfam. Prot(‘eae- -

(e) Proteaceae.

(2) E. Australia
( I ) Tasmania
(1) Tasmania
(7) New Caledonia
(1) Madagascar
( 1 ) Tasmania
(1) New Caledonia
(62) Australia and New' Zealand
( I ) South Africa

Aulax*

(3) 8. Africa

Lencadpyulron *

(70) 8. Africa

Protea *

(75) 8. and trop. Afr.

Lencosperuemu *

(40) 8. Africa

Faurea*

(10) Africa and Madagasc

Serruria *

(50) 8. Africa

Mimetes*

(15) 8. Africa

Sorocephalus*

(10) 8. Africa

Parano7tuis*

(12) 8. Africa

Orotha7unnf>*

8. Africa

DiastelUP

8. Afi'ica

S pa f alia *

(25) 8. Africa

Spatallopsifi*

(5) 8. Africa

Petrophila

(35) Austr. (W'.A.)

Isopogon

(30) Austr. (W.A.)

Adenanthofi

(20) S.W. Austr.

Stirlingia

(5) S.W. Austr.

Synaphaea

(8) S.W. Austr.

Conospermutn

(35) Australia

FrayikUtndia

(2) S.W. Austr.

The Vegetation of Western Australia.

xix.

Subfam. (Jrevilloideae- — •

Darlingia*

(1)

E. Austr.

BucMngha^nia*

(I)

Ph Austr.

Greinllea

(180)

Austr.

Strangea

(3)

Austr.

Carnarvonia'^

(1)

E. Austr.

Hakea

(100)

Austr.

Orites*-

(6)

Ph Austr.

Helicea*

(40)

Japan, Tndo-Malaya, E. Austr.

Xylomelum

(4)

Austr.

LamheHia

(S)

Austr.

Roupala^

(40)

South America and Ncnv Caledonia

Panop sis*

(8)

trop. America

Macadamia*

(5)

E. Austr.

HicksbeacMa*

(1)

E. Austr.

Kermadecia*

(5)

P]. Austr. and New Caledonia

Ouevina*

(1)

Cdiile

Euplassa*

(8)

tro]'). America

Embothrium*

(•>)

Chilean Andes and PT Austr.

Telopea*

(3)

K. Austr. and Tasmania

Lomatia*

(10)

p]. Austr., Tasmania, and Chile

Knightia*

(3)

N. Zeal, and N. Caledonia

Cardwellia*

(1)

E. Austr.

Ste7iocarpus

(15)

N. Caledonia and trop. Austr.

Banksia

(50)

Austr.

Dryandra

(52)

S.W. Australia

Summary (54 genera).

21 genera endemic in Australia.

14 genera endemic in S. Africa and Madagascar.

3 genera endemic in America.

2 genera endemic in Tasmania.

2 genera endemic in New Caledonia.

1 common to S. America and N. Caledonia.

2 common to S. America and E. Australia.

2 common to New Caledonia and E. Australia.

1 common to Australia and New Zealand.

J common to Japan, IndoAIalaya, and E. Austr.

1 genus endemic in Madagascar.

The preceding tables are characteristic of the groups within the Ant-
arctic Element. They show affinities between the three great southern land
masses, with Madagascar, New Caledonia, and New Zealand as intermediate
j^oints. It is interesting to find that, in general, the links between Australia
and South America are usually stronger than those between Australia and
South Africa. These will be referred to again under the Australian Element.
Amongst the Restionaceae there is a high degree of endemism both in the
Australian and South African groups, with one conjunctive genus for America
and one for South Africa. The Centrolepidaceae are more strictly Antarctic,
and the link here is between South America and Australia. The same holds
good for the Epacridaceae and Stylidiaceae, whieli are not represented in
South Africa, the former without conjunctive genera, the latter with two.
Amongst the Proteaceae however, there are powerful links with South America,
in the Orevilloideae, with two conjunctive genera ; while amongst the Proteeae

XX.

Charles Austin Gardner.

the development in South Africa is even greater than in Australia, and there
are again no conjunctive genera. In all cases in the Proteaceae, where a
genus exists in South-Western Australia, it is more powerfully developed
there than elsewhere.

At the same time, there are several genera common to both Australia
and South Africa, some of which will be discussed under the Palaeotropic
Element, but the following are genera confined to the countries mentioned : —

Chrysithrix

Tetraria

Wurmhaea

Biilbine

Caesia

Podocoma

Trichocline

South Africa and S.W. Australia.

.... South Africa and S.W. Australia.

.... South Africa and S.W. Australia.

.... South Africa and temperate Australia.

South Africa and temperate Australia.
.... Australia and America.

.... S.W. Australia and S. America.

These are but a few examples of the Antarctic Element, and when floral
statistics have been completed for Western Australia, further additions will
probably be made. Sufficient evidence has, however, been advanced to
indicate the close relationships which exifet between certain southern groups.

2. THE PALAEOTROPIC ELEMENT.

The Indo-Melanesian Element is expressed by those plants which are
common to Australia, Indo-Malaya, and Melanesia. In Australia they are
most in evidence along the Northern Australian littoral, especially in the
high rainfall districts of Queensland, and extending through the rain-forest
into New South Wales. The Element is also fairly well indicated in Western
Australia in the Kimberley district, especially in the littoral, the riverain
forest, and the monsoon woodland. The conditions governing its migration
are the requirements of a megathermic flora, and, while it is most strongly
represented in Western Australia in the Kimberley district, it has migrated
fairly extensively along the littoral tracts. This must have taken place
at a time ■when the climatic conditions were very different from those of to-day.
The Element has not only strongly established itself in Kimberley, but its
residuals may be found fairly well represented to-day in the De Grey River
district, and the Hamersley plateau. At one time it extended as far south as
the South-West, for it has not only imdoubtedly exerted an influence on the
flora of the South-West, there are today certain representatives of this Ele-
ment as far south as the karri forest. There can, indeed, be no other ex-
planation for the occurrence of such plants as Cartonema, Dioscorea, and
Clematocissus in regions between Geraldton and Perth. At the same time, such
a direct line of commmiication would explain certain northern migrations of
plants that are typically southern, e.g., Byhlis, certain Droserae and Jacksonia,
to the tropics. Finally, if further evidence is wanted, one has but to consider
the flora of the Pilbarra district — the region between the De Grey and Ash-
burton rivers — ^which is today an ‘‘island” supporting the impoverished
I'esiduals of the Indo-Melanesian Element under conditions which are generally
imsuitable, either as lithophytes or species of the declivities. Examples such
as Livistona, Astrotriclia, Owenia, and several herbaceous species, also the
few endemics, such as Elachnanthera, as megathermic plants of mesophytic
or hygrophytic derivation, are evidence of a former pluvial area which must
have been of importance in bridging the line of communication between the
north and south.

The Vegetation of Western Australia.

XXI.

A list of the gi'oups which comprise the Indo-Melanesian Element would
be beyond the scope of this treatise. It is sufficient to state that it includes
the Palaeotropic Rhizophoraceae, many Leguminosae and Gramineae, Santa-
laceae, Lauraceae, Menispermaceae, Ulmaceae, Moraceae, Burseraeeae, Melia-
ceae, Euphorbiaceae, Anacardiaceae, Celastraceae, Sapindaceae, Rhamnaceae,
Malvaceae, Tiliaceae, many of the Parietales, Myrtiflorae-Myrtoideae, the
Primulales, and a host of tropical Sympetalae.

While we postulate a land connection for the greater number of plants
comprising the Palaeotropic Element, it must be remembered that a number
have undergone an oceanic migration. Such plants as the Rhizophoraceae,
Terminalia, Carapa, Thespesia, Barringtonia, and Camptostemon being ex-
amples, as well as the maritime Spinifex and certain dune species. They
remain principally as littoral plants.

In addition to those groups or species which we regard as belonging
essentially to the Palaeotropic Element, there are several groups which, while
perhaps or palaeotropic origin, have become so modified, or have undergone
such an extensive development, that they are no longer recognisable as typical
examples. Indeed, in some instances they have assumed such importance in
Australia that they have been regarded by some authors as typifying the
Australian Element. I propose to consider, by way of illustration, one Order,
tlie Leguminosae.

In the first family of this Order is the genus Acacia, This genus, whicli
is represented in the tropics of both the east and west hemispheres, com-
prises over 500 species, of which 400 are indigenous to Australia. Of the
six Sections, four are represented in Australia (all being in Western Aus-
tralia), the remaining two being found in America, Africa, and Asia. The
most primitive form of Acacia is that which possesses compound leaves.
Acacia Farnesiana, which may be regarded as an archetype, inhabits the
tropics of both the old and new worlds, extending from America, through
Africa, Arabia, and India, into northern Australia. In Western Australia it
extends as far south as the Murchison River, while in eastern Australia it
occurs as far south as the interior of New South Wales. Within the high
summer rainfall areas, e.g., in Kimberley, it is of fairly general distribution
as a shrub of the savannahs, but south of the Fitzroy River it is confined to
the alluvial soil of watercourses and depressions.

The remaining species of the Section Gumniferae in Australia are all
endemic and of tropical distribution as far as Western Australia is concerned.
Acacia Farnesiana alone of the Bipinnatae connects the tropical species of
this genus with the temperate Bipinnatae in south-w'estern Australia. Acacia
pulchella, the most widespread of the latter, extends northwards to the south-
ern limits of Acacia Farnesiana and southwards to the forest areas where it
connects with the remainder of the species of its Section, the Pulchellae, which
number twelve, and are of considerable importance in the high rainfall forest
area of the south-west. Apart therefore, from a northern gi’oup consisting
of the Gumniferae and the single representative of the Botryocephalae (A,
pachyphloia), and a southern group in which there are 12 species of the Pul-
chellae, the two being connected by the riverain A, Farnesiana, there are no
other leafy Acacias in Western Australia. In eastern Australia a similar
development may be noted, Acacia Farnesiana continuing the line of de-
velopment into New South Wales, while a south-eastern group (Botryocephalae)
extends from the north to Victoria and South Australia.

XXll

Charles Austin Gardner.

From this original stock, a type of Acacia has originated which is
characterised by the production of phyllodes, the leaf being represented by the
simple flattened leaf-like axis of the compound leaf. Its ontogeny may be
observed in any species of this type, and transitions between the two may be
observed in Acacm insolita, a species of the eastern borders of the jarrah
forest, w^hich normally possesses both leaves and phyllodes. This large
grou }5 (the Phyllodineae), numbering over 400 species, is, with the exception
of a few Papuan and Pacific species, confined to Australia, the Western Aus-
tralian species numbering over 300. They are found wdthin every formation
throughout the country, assuming remarkable diversity in form and varying
from complete aphylly to large leaf-like phyllodes. Their highly developed
floral economy, and the adaptation of a leaf-stalk or even a stem fimctioning
as a leaf, has enabled them to withstand extreme aridity of environment,
and thus they have penetrated into the arid heart of the continent, developing
an epharmony which leads to complete aphylly.

While this epharmonic development is in most examples progressively
manifested as w'e travel from w^etter to drier areas, there are w^hat appear
to be retrograde divergences in some instances. For example. Acacia alata
and A. cxtcnsa, both of w'hich inhabit wet or marshy spots in the south-western
forests, exhibit an apparent morphological xerophily which is misleading,
for histologically Acacia alata at least, is a mesophyte. It lives only under
conditions of shade and moisture wfiiich are the ideal conditions under which
the leafy Acaciae find their true environment ; the phyllodineous species, on
the other hand, have attained to their highest development under arid con-
ditions, or at least, seasonal droTight. Australia, and particularly Western
Australia, is the richest in species of Acacia of any part of the earth, but the
genus cannot be considered as belonging to any but a tropical element.

What is true of Acacia is ajjparently also true of the rest of the Mimosaceae
and the Caesalpiniaceae. In this connection it is interesting to consider the
range of Albizzta, indigenous to tropical Asia, Africa, and Australia, fhe
Western Australian representatives are confined to the trojiical north with
the single exception of Albizzia distachya, confined to the south-w^est. Of
the Caesalpiniaceae, both Petalostyles and Labichea are endemic in Australia,
and are closely related to Cassia, from which they have probably been evolved ;
Petalostyles remaining megathermic, while Labichea extends from Queensland
to south-wustern Australia, but without a continuous distribution between
the Murchison River and the Northern Territory. Cassia, on tlie other
hand, which w'e must regard as a tropical element, has migrated far to the
south, but has not extended into the wet south-wust, and its real liome in
Australia in the savannah and the dry interior, where, by the provision of a
dense indumentum of felt or hair, it has, with practically no structural modi-
fication, iDenetrated to the desert.

Space does not permit me to deal at any length with the third family of
the Leguminosae — the Papilionaceae, and the family is so large, and of such
cosmopolitan distribution, that it is difficult to trace with any exactness its
possible migrations. It is w'orth remembering, however, that every tribe
of this important family is represented in America, which country appears to
be the main centre of distribution for the family. The Dalbergieae, Sophoreae,
Hedysareae, Swartzieae, Galegeae, and Phaseoleae are all most richly de-
veloped in America, five of them being found in Asia and Australia. Of the
remaining five tribes, the Vicieae, Trifoleae, and Loteae arc more cosmo-
politan in distribution, but mainly of northern development, while in the

The Vegetation of Western Australia.

XXlll.

last two — the Genisteae and Podalyrieae — there is a marked southern de-
velopment. The following list gives the distribution of these two tribes : —

Genisteae. Podalyrieae.

South Africa

19 genera

Australia ....

20

genera

North Africa and

8 genera

North America and

3

genera

Mediterranean

Siberia

Australia ...

7 genera

Himalayan

2

genera

Warm regions

2 genera

Mediterranean

1

genus

Indo-Malayan

2 genera

South Africa

1

genus

N. Hemisphere

(mesothermic) ....

1 genus

South America

1 genus

It is suggested that both tribes migrated southwards from the northern
hemisphere, and that the Axistralian Podalyrieae, like the South African
Genisteae, have formed their own centres of distribution. It will be noticed
that the Podalyrieae are absent from both New Zealand and South America,
and that botli have a number of localised endemics in two centres — the Genis-
teae in the Cape Peninsula of South Africa, and the Podalyrieae in southern
Australia.

Finally there is a relationship between the flora of Madagascar and Aus-
tralia that cannot be overlooked. There is, in the first ])la(5e, the remarkable
distribution of the Baobab (Adansonia). This genus is restricted to the
African savannahs, Madagascar, and that portion of Australia wliich lies
between the Fitzroy and Victoria rivers. In the second place there are those
genera which are entirely restricted to Madagascar and Australia — -and p^u'-
ticularly well developed in Western Australia- and Kermidrenia

(Sterculiaceae), and Diplopeltis (Sapindaceae). No attempt is made to
explain this rem^irkable distribution.

3. THE AUSTRALIAN ELEMENT.

Botanists and zoologists luiN'e rep(‘atedly referred to an Australian Ifli'inent.
Hooker, speaking of the Australian Flora, states : “ It contains more genera
and species peculiar to its own areas, and fewer j)lants belonging to other
parts of the world, than any country of equal extent. About two-fifths of its
genera, and upwards of seven-eights of its species aix^ entirtdy confined to
Australia. On the other hand, if, disregarding the pt'culiarities of the flora,
I (ionipaix' its elements with those of the floras of similarly situated larg(' areas
of land, or with that of the whole globe, I find that there is so great an agree-
ment b(>twoen these? that it is impossible to i*(?gard the Australian vegetation
in any other light than as forming a peculiar but not tu) aberrant or anomalous
botanical ])rovince of the existing Vegetal)le Kingdom ; that with only two
small exceptions, the Australian families are also found in other countries ;
that most of those most widely diffused in Australia are such as are also the most
wid(fly distributed over the glob(‘ ; and that Australia wants no known family
of general distribution. Tliat the largo families and gc^nera which, though
not absolutely restricted to Australia, are then* very abundant in sp(‘cios and
rarc^ elsewhere, an<l for which 1 shall hence adopt the t(‘rm Australian, stand
in very close relationship to groups of plants which ar<* widely spn'ad over the
glob(? {as Kpacridaceae to Fricaceae ; Good(‘uiac(‘a(? to (<ampanula(?eae ;
Styli(liac(‘ae to LolH'liaceae : ('asuarinaceao to Myrica(H'ao). Turning again

to other countries vhich are remarkable for tlio peculiarity of tlu?ir vegetation,

XXIV.

CuARi.Ks Austin Gardner.

1 find that South America contains many more* peculiar families than Australia,
and South Africa about as many.” Tlie definition given by Diels for the Aus-
tralian Element has already been quoted (p. 3), and 1 ha\'e already indicated
that some of the groups included under his definition do not belong to this
element.

On the other hancl, there are certain families and groups tliat are either
entirely Australian, or almost entirely so, vitli every genus represented in Aus-
tralia, and but few external species. Apart from the strictly endemic groups
it is difficult to deline with exactitude the Australian Element, or to di\*orce
it entirely from the Antarctic, or the Palaeotropic Element. To illustrate this
difficulty we may take the sub-family Leptospermoideae of the ^Myrtaceae.

MYKTACEAE UEPTOSPERMOmEAE.

Osho^'nia"^

Back'housia*

Metrosideros*

Bpe?'7nolepis*

Lysicarpvs*

Cloezia*

Tepualia*

jSyncarpia*

Xanthoste^non

Fleur ocaly pin

Tristania

Angophoi'a'^

Eucalyptus

Leptospennum
Agonis
Kurizea
Callistemon
M elaleuca

Lamai'chea

Coiiothatmms

Eremaea

Ph ym at 0C07 pus

Regelia

Beaufortia

Balaustio7i

Baeckca

A startea

Hypocahjmma

(a) Leptospermeae.

(1) N. Australia

(5) N. Australia

(20) S. Africa, Indo-Malaya, New Zealand, Aus-
tralia, Sunda Is.

(2) New Caledonia

(1) N. Australia

(6) New Caledonia

(1) Chile

(2) E. Australia

(18) trop. Austr., N. Caled. .... .. . 1

(1) New Caledonia
(22) Malaya, N. & E.

Australia, N. Caled .... .... 3

(5) E. Australia

(500) Australia, Malaya, Celebes, Philip-
pines .... .... .... .... 150

(25) Australia, Malaya, N. Zeal. .... 10

(13) S.W. Australia .... .... .... 13

(18) temp. Australia .... .... .... 12

(12) temp. Australia ... .... .... 2

(120) Australia — one sp. extending to N.

Caled., Malaya, and India .... 100

(1) S.W. Australia .... .... I

(3) S.W. Australia .... .... .... 3

(7) S.W. Australia .... .... .... 7

(2) S.W. Australia .... .... .... 2

(3) S.W. Australia .... .... .... 3

(15) S.W. Australia .... .... .... 15

(2) S.W. Australia .... .... .... 2

(00) Australia, Malaya, China .... .... 48

(5) Australia .... .... .... .... 5

(18) S.W. Australia .... .... .... 18

The Vegetation op Western Australia.

XXV.

(b) Chamaelaucieae.

Lhotzkya

(10) temp. Australia

7

Calythrix

(40) Australia

.... 37

Calythropsis

(1) S.W. Australia

1

Homalocalyx*

(2) N.E. Australia

Micromyrtus

(13) Australia

.... 12

Wehlia

(5) S.W. Australia

5

Pileanthus

(3) W. Australia ....

3

Chamaelaucium

(12) S.W. Australia

. 12

Thryptomene

(22) temp. Australia

.... 21

Actinodium

(1) S."W. Australia

1

Darwinia

(31) temp. Australia

.... 27

Homoranthus*

(2) E. Australia

V erticordia

(47) W. and N. Australia

.... 47

Summary.

Sub-family Leptospermoicleae .... 41 genera.

Australia .... .... .... 31 genera (15 endemic in S.W. Austr.)

Austro-Malaya, extending to
India, China, New Zealand,
and 1 (Metrosideros) S. Africa 6 genera

New Caledonia ....

3

genera

Chile

1

genus

Sub-family Myrtoideae

.... 32

genera

America ....

.... 22

genera (mainly S. America)

New Caledonia ....

3

genera

Indo-Malaya

2

genera

Malaya- Australia

3

genera

Australia ....

1

genus

Tropical Africa ....

1

genus

' Tropics (general)

1

genus

In the above, and succeeding table, the first column of figures in paren-
thesis represents the total number of species in the genus ; the second indicates
the number found in Western Australia. Names indicated by asterisks are
those of genera not occurring in Western Australia.

It will be seen that of the two sub-families of the Myrtaceae, the Lepto-
spermoideae is predominantly Australian, but that three genera are restricted
to New Caledonia, while one is South American. Of the Australian genera,
half are endemic in South-Western Australia. In the sub-family Myrtoideae
(those with baccate fruits), the position is quite otherwise, the majority of
genera being endemic in America. The Chamaelaucieae, a tribe of the Lepto-
spermeae, is strictly Australian, and this group alone, unless we include the
nine endemic genera of the Leptospermeae, can be defined as an Australian
Element. The southern, or Antarctic influence is expressed by Metrosideros
and Tepualia. In any case, we see once more that the Australian Element
is most richly developed in South-Western Australia.

XN \ 1.

Chari. K s Austin (Gardner.

An almost parallel ease is exhibited by the Goodeniaceae, but here eacli

genus is represented in Australia : —

VelUia (18) Australia 10

Symphyobasifi (2) Australia .... .... .... .... 2

Goodenia (*01) Australia 56

Calogyne (5) Australia (one extending to the Philip-
pines) .... 3

Leschenaultia (19) Australia .... .... .... 17

Anthotiuni (2) S.W. Australia

Selliera (2) One endemic in S.W. Austr., the

other extending from Victoria and
Tasmania to N. Zealand and Chile I
Pentaptilon (1) S.W. Australia .... ■■■■ 1

Catosperma (1) N. Australia 1

Diaspasis (1) S.W. Australia .... .... .... 1

Scaevola (83) One trojiics generally ; 22 Pacific and

Melanesian (excl. Australia) ; Austr. 50
Verreauxia (3) S.W. Australia .... . .. . .. 3

Danipiera (59) Australia .... .... ... 48

This family represents the true Australian Element. It agrees with
the definition given by Diels, for each genus is represented in Australia, only
three having extraterritorial representatives. The species connecting S.E.
Australia with Chile is again interesting. It will be observed that each genus
is represented in Western Australia. Apart from Selliera and Calogyne,
the migrant species are all species of Scaevola, entirely or chiefly of maritime
distribution.

In addition to the three endemic families — the Tremandraceae, Byblida-
ceae, and Cephalotaceae, there are a number of groups with a rank less than that
of family which are endemic. Examples are : —

Liliaceae-Xanthorrhoeae

Haemodoraceae-Conostylidiae

Pittosporaceae-Billardiereae

Rutaceae-Eriastemoninae-Nematolepidiuae-Diplolaeninae.

fkiphorbiaceae-Stenolobeao

Myrtaceae-Chamaelaucieao

Verbenaceae-Chloanthoideae

Labiatae-Prostantheroideae

Oompositae-Angianthinae

The number of genera is too large for inclusion here, but examples an' to
bo found in many families. The total number has not been accurately com-
puted.

In every case, the Australian Element is most richly developed within
the South-West Province ; its real home today is the large triangular-crescentic
area bord(>r('d by the coastline from Shark I3ay to Israelite Bay, and extending
inland to the 175 mm. (7 in.) winter isoliyct. Within this area the autoch-
thonian flora finds its highest expression in the sandy areas, either on the sand
iK'aths, or in the sandy swamplands of tlie south-western littoral. W hile many
of its groups are strictly endemic within the area, others are found in Eastern
Australia, esj)ecially in Southern Quet'iisland and South Australia, where
tluiy occur as psammophytes. They arc foum^l also in the littoral region
of South Australia, whore their occurrence can be explained by littoral migra-
tion, but their occurnnce in soutlu'rn Queensland and parts of New South Wales
is more difficult of explanation.

Tut’, Vkgetation op Western* Austijalia.

XXVll.

These eastern species and groups bear the same relationship to tliose
of South-Western Australia that the groups in South Africa and SoutJi America
bear to the Australian ingredients of the Antarctic Klement.

Whether South-Western Australia w'as the cradle of the Australian Ele-
ment, or whether it is here alone that it has maintained its floristic stability,
is a ({uestion which has not yet been decided. Wlien we consider tlie liigh de-
gree of endemism within the area, and its ec^ological stability, one is inclined
to the foi-mer view, but, as Diels has pointed out, the occurrence of the \-ast
epicontinental sea of Cretaceous times must hav(‘ caus('d a division of what may
originally have been an ancient pan-Australian flora, and the present distri-
bution of such genera as Borya, Petrophila , Jsopogon, Grevillea, and Banksia,
to enumerate a few only, lends a.dehnite support for tlie tlumy of the previous
existence of a pan-Australian Element. ]n any cas(‘, it is in South-Western
Australia alone that the true Australian flora is today most richly developed
within a climatic and geologically distinct entity, in an almost perfect state
of efjuilibrium which is disturbed only by tlu' gradual process of climatic
desiccation in operation everywhere. It is true that a disturbing influence
has to some extent been at work since the ach-ent of settlement : agriculture,
with grazing, fire, and the plough, has done much to bring about important
local changes in the country’s physiognomy, but tlu* work of man, and the plants
which he has brought wdth him, including those aliens which we regard as weeds,
although of great importance in man’s particular economy have, in the main,
proved quite ineffective as a permanent invading force when subjected to
competition under natural conditions with the indigenous vegetation.

III.—PHYSIOGRAPHY.

Western Australia consists almost entirely of a low" plateau with an
average elevation of between 1,000 and 1,500 feet. Between Point Culver
and Eyre on the south coast, between the King Leopold Range and Van-
sittart Bay in the Kimberley district, and in a few" other places, this plateau
extends to the coast. In other places, e.cj., between Eyre and Eucia, and
along the western coast betw'een the Moore and Vasse rivers, the plateau
descends rather abruptly to a narrow littoral plain. In the north-west, be-
tween the Murchison River and Broome, the plateau descends gradually to
the coast, or the escarpment is far inland, exc^ept in the Pilbarra district,
where the Hamersley massif extends almost to the coast at Vlaming Head,
and again in the vicinity of Roebourne. While little is knowm of the contours
in the vicinity of the Eighty-mile Beach, the Gascoyne, Ashburton, and Eortes-
cue rivers in their lower courses flow* over broad alluvial flood-plains.

The surface of the plateau is mostly flat or gently undulating. According
to Jutson, the main plateau represents what is probably a vast uplifted pene-
plain now in a state of arid erosion, bearing the residuals of a former higher
plateau which has suffered very considerable erosion. There are no mountains
in the sense of large folded and not much denuded systems, and the high
points w"hich rise above the existing plateau are in the main either residuals
of the former higher plateau erosion, or fault blocks. Tlie highest elevations
are found in the Hamersley Range, w"hcre Mount Bruce attains 4,024 feet.
In the Kimberley district, the King Leopold Range rises to nearly 2,000 feet.
This range is the escarpment of the Hann ])lateau which extends almost everv-
where to the coast to the north and west, and is boun{ted on the east by the
Durack Range. It culmina,tes in Mount Hann (2,800 feet). In the south-
west, the main physical features are tlie Stirling Range — a system of isolated
peaks, of which Coyanarrup is the highest (3,(>40 feet) — while to the south
is the Porongorup Range, and parallel with the soutli coast is a system of

XXVlll.

Charles Austin Gardner.

isolated hills extending from Mount Manypeak through the Barren Ranges
to the Eyre Range and the Russell Range near Israelite Bay. The Darling
escarpment averages about 1,000 feet in height, the highest point being Mount
William (1,689 feet). This escarpment extends from near the south coast
as far north as the Irwin River, and possesses two subsidiary spurs, one ex-
tending to the north-west from Gingin, the other, below Bunbury (known
as the Roe Range) links the main escarpment with the Dunsborough- Augusta
escarpment between Cape Naturaliste and Cape Leeuwin.

Most of the drainage is into the sea. While a number of streams are
short and originate near the margin of the plateau, others, like the Fortescue,
Ashburton, and Gascoyne, are of considerable length, and originate on the
plateau. The rivers of Kimberley rise in the Hann plateau ; with the ex-
ception of the Fitzroy and Lennard rivers, they are perennial, at least in the
plateau area, and they have, together with their tributary streams, eroded
narrow valleys or canyons in the soft sandstone. They appear to be yoimg
rivers, and several of the smaller tributaries have in places tunnelled through
the soft rocks. Most of them rise in the comparatively high rainfall area
near Mount Hann.

South of the Fitzroy River, as far as the Avon, the streams are inter-
mittent, and flow only in times of heavy rain. They are, in fact, merely
flood channels which carry water in times of flood. They may remain as
dry watercourses, containing only a few shallow pools, for several consecutive
years, or, on the other hand, they may flow twice or more during the same
year. They traverse low flood plains in their low'er courses, and a character-
istic of the larger streams is the formation of subsidiary channels of “ billa-
bongs.”

The Swan River marks the commencement of those streams which are
more permanent and have eroded broad valleys. They extend southwards
to the Denmark and Kalgan rivers. These rivers flow as a rule throughout
tlie winter months, but in the summer are much reduced, sometimes to chains
of pools, or in the lower south-west, as for examples the Warren and Frankland
rivers, they may be permanent. Some of them, such as the Avon, drain
larger watersheds than appear to be indicated on the map as their defined
courses, and it is probable that the Avon River once drained a very extensive
area, since the chain of salt pans and channels which enters the river at Lake
Mears, can be more or less continuously traced through Lake Brown to Lake
Ballard. It is only during exceptional seasons that this system flows to the
sea from above Hine’s Hill, thus accounting for tlie salt-j^an system of the
upper course.

Apart from those streams which find an outlet to the sea, there are a
number of watercourses which very rarely carry rimning water through more
than a limited portion of their channels, emptying themselves into extensive
flats or clay-pans. Some of these clay-pans are termed "salt lakes.” \\ith
the eva]joration of the water in these depressions substances such as salt and
gypsum are deposited. Many of these so-called ‘'lakes” may be of con-
siderable extent, such as Lake Moore and Lake Carnegie. They are either
fpiitc isolated, or they are constellated over the interior in a manner that
suggests that they once formed old river beds forming part of an ancient river
system. The largest of these ancient systems is that which extends from
Lake Nabberu to Goddard’s Creek near Zanthus. Its ancient course once
entered the sea near Twilight Cove, and it constitutes the largest watershed
in Western Australia.

The physical features described are important because of their effect
ujDon the flora, and upon the climate. The river systems are important be-

The Vegetation op Western Australia.

XXIX.

cause not only do they support their cliaracteristic vegetation, but because
they have been of importance in plant migrations. The salt pans provide
their own halophytic associations.

IV.— GEOLOGY AND SOILS.

Apart from the climatic factor, the edaphic factor is the most important
in j^roviding the particular requirements for individual plants, and in de-
termining the characteristics of plant communities. The nature of the soil
is dej:)endent upon the original rock formations which are fundamental in
determining soil formation and soil characteristics.

The geological fundament of Western Australia is a Precambrian com-
plex, consisting mainly of metamorphic rocks and granite. The granite
rocks are exposed in many parts of the territory, particularly in the south, and
especially in the Darling and Porongorup Ranges, and parts of the interior
of the plateau where they usually occur in the form of convex bosses or tors.
Wherever the granite is exposed, or lies close to the surface, it exerts an in-
fluence on the plant life, either because of the immaturity of the derived soil,
or by reason of the water which it collects. Especially in the arid interior,
the flora of the granite rocks stands in marked contrast to that of the sur-
rounding country.

Superposed on these basement rocks are many sedimentary series. Of
these, the oldest is that knovm as the Nullagine. The rocks of this series are
freely exposed in the Kimberley and Pilbarra districts, where they form the
most prominent physical features of these areas. In the Kimberley district,
on the Hann Plateau, and on the Antrim Plateau, the base of the Cambrian
is composed of a series of basalt “flow's.” The occurrence of basalt in the
surface soil determines a distinctive type of savannah or savannah-w'oodland
in which the species of Themeda, the “ Kangaroo-grass,” is dominant, while
the indicative prevailing tree is Eucalyptus Spenceriana. The “Nullagine”
rocks, on the other hand, support on their derived soils a type of steppe or
impoverished savannah in w'hich Triodia takes a prominent part in the physi-
ognomy.

The Devonian strata, typified by the Oscar and Napier Ranges, are im-
portant because of the more richly varied flora w'hich they support. In the
Napier Range we find Melia common, and several species not found in the
neighbouring soil types. The number of endemics here is interesting.

The Permian rocks of the Gascoyne-Ashburton divide appear to be im-
i:»ortant in more respects than one. The area of tlieir occurrence here co-
incides with that part of the Mulga-bush formation in wdiich the true Mulga
(Acacia aneura) is either rare, or entirely absent, and the general physiognomy
is more reminiscent of the Pindan country, a type w'hich is common along
the low'er Eitzroy River, and paits of Dampier Land. The hard-pan soil,
so typical of the true Mulga formation, appears to bo (juite absent from these
areas.

The rocks of the Kainozoic Era are associated wflth characteristic featiux's
o^ the vegetation. They occm as sedimentary formations in many parts
of the w'estern and southern littoral tracts, extending as far north as Vlaming
Head, and on the south coast as far as Eucla. Between King George’s Sound
and the Phillips River they occur in the form of siliceous sediments, and
sup])ort a well-dif£erentiated flora ricli in many endemic IMyrtaceae and
Proteaceae, mostly in the form of heath or low thicket formations ; finally,
in a calcareous form they provide the basis for the phytogeographically distinct
area of low salsolaceous shrub-steppe that characterises the Nullarbor Plain.

XXX.

Charlks Austin Gardner.

The superficial deposit known as laterite by Western Australian geologists,
is extensively developed on the Darling Plateau, and occurs in many parts
of the southern interior, either capping the higher elevations in the form of a
more or less continuous cuirass, or superficial accumulations of pebble-like
lateritic nodules with varying amounts of clay and sand. Laterite also occurs
near the northern edge of the Hann Plateau between Admiralty Gulf and
the Couchman Range in Kimberley, associated there with palm woodlands
and a type of sclerophyllous woodland. Laterite also provides the edaphic
requirement of the jarrah forest. In the interior it is frequently associated
with an impoverished sclerophyllous association in areas of low rainfall,
typified by the “ Wodjii ” type of country, certain Casuarina thicket associa-
tions, and sometimes with small dwarf woodlands of Callitris. The lateritic
areas are usually the most deficient in grass, but surprisingly rich in Goodeniaceae.

Finally, mention should be made of the sand which is so characteristic
of large areas of the south-west, in the far interior, and in the littoral regions
between the Gascoyne and Ashburton rivers, and again between the De Grey
and Fitzroy rivers. Within the south-west, on the plateau, it is often difficult
to divorce the sand from the laterite as a distinct formation, since laterite so
frequently vinderlies the sand. But in any case, it is on the sand that the
shrub-heaths hold undisputed sway, whether the sand be the loose detritus
of the coastal ]dain, the humus sand of tlie southern littoral, or the more
compact yellow or red sand of the interior. Even the sand of the desert
supports psammojfiiytes with structural peculiarities, enabling them not
only to thrive in a hostile environment, but sufficiently plastic to create new
forms. Finally, it is in the sand that the south-west elements have made
their furthest advance into the interior.

V. CLIMATE.

The position of Western Australia with reference to its range in latitude
(almost equally tropical and extratropical), its low elevation and uniform
topography, together with the fact that it lies on the western side of the con-
tinent, and is, therefore, subjected to the desiccating effect of the south-easterly
trade-winds, are facts of fundamental importance from the climatic standpoint.
Tliese facts account for the different climatic systems, and for the great arid
mi<ldle region wliich represents tlie western prolongation of the desert of
Onti-al Australia.

Three climatic zones may be recognised : a northern area of summer

rainfall of a monsoonal character, with a cool dr^^ season ; a south-western
area of winter rainfall with a period of summer drought, and a vast central
ai't^a of low and unreliable rainfall of no marked periodicity, depending en-
tirely upon extensions of the climatic systems which dominate the northern
and southern areas.

1. THE CLIMATIC CYCLE.

In order to understand the three climatic areas it is necessary to give
a bri(T account of the operation of the systems.

In January and February, the thermal equator lies over the north, where
high temperatures are experiencetl, especially over the Pilbara and Hamersley
districts, which at this time are amongst the hottest regions of the earth’s
surface. The intense heat causes an inflow of cooler air from the tropical ocean
—a monsoonal system which brings rain from the north-west. A lag ” of
from four to six weeks which occ\irs between the atmospheric movements and
the apparent solar movement accounts for the retardation of the season
which normally extends from December to March. There are two pluvial
foci : one where the Hann Plateau extends to the coast between the King

Leopold Range and Admiralty Gulf, where the seasonal precipitation exceeds

The Vegetatiox of Western Australia.

XXXI.

1,110 mm. (4,300 points), and a secondary focus of much less importance
in the Hamersley Range. The absence of meteorological stations on the
Hamersley Plateau leaves us in ignorance regarding the local climatic con-
ditions, but the area experiences an increase in rainfall compared with the
lower country which surrounds it. From both of these two focal points there
is a gradual diminution in the rainfall. The rainfall decreases sharply between
Camden Sound and the southern and eastern margins of the Harm Plateau,
being about 490 mm. (1,934 points) at Broome, and 567 mm. (2,229 points)
at Wyndham, while at Wollal it falls to 249 mm. (978 points). The seasonal
isohyets rise to 275 mm. (1,090 points) at Whim Creek and Tambrey, at the
northern end of the Hamersley Range, and are doubtless higher within the
Range itself. Over the whole of the north the rainfall is very low between
the months of April and November, being everywhere less than four inches,
and in places below one inch.

A climatic feature of some importance, even if not of any marked seasonal
reliability, is the occurrence, usually between the months of January and April,
of tropical hurricanes which, bom in the Timor Sea, sweep in a south-westerly
direction over the sea until they reach the approximate latitude of 18 degrees
S., when they curve in a southerly or south-easterly direction and encounter
the coast, usually between Broome and Onslow. These violent disturbances
may expend their energy soon after reaching the land, or they may continue
inland, bringing with them heavy rains, often extending as far south as the
latitude of Wiluna, and occasionally to the Nullarbor Plain. They are quite
unreliable in their incidence, sometimes occurring twice or more in a given vear,
at other times they may be entirely wanting for two or more years. Their
importance as the principal rain-bringing systems of the great central area,
is considerable, smce they determine the habitat of the grassland and steppe
of the arid region.

While this northern summer monsoonal system is in operation, the south-
ern half of Western Australia lies under the influence of high-pressure systems
and summer drought, and except for occasional thunderstorms wdiich are either
<lirectly or indirectly influenced by the low’-pressure systems of the north, little
or no rain falls except in the extreme south-west and southern littoral where
the karri forest receives 10 inches of rain, and the entire south coast five
inches during this period, due to northw^ard surgings of the systems carried along
by the “ roaring forties ” at all seasons in the Southern Ocean.

As the apparent northern motion of the sun tow’ards tlie Tropic of Cancer
progresses, so the thermal equator shifts to the north a\\ay from Australia,
and the tropical area lies under a system of laigh-pressure and drouglit which
continues from April until November or December. This northern “ shift ”
of the atmospheric systems causes a similar northern shift of the so-called
“ Antarctic ” systems, bringing into operation the cycle of whiter precipitations.
Tlie winter rainfall cycle usually commences in May, either early or late in tlie
month. Sometimes it commences in April, and is then more or less connected
with the late cyclonic systems of the north-west (Ashburton-Fortescue) areas.
If, as sometimes happens, a powerful southern system should coalesce w'ith a
nortlK'rn active low^-pressure system, the result is a general rainfall o\’er the
entire west coast, usually regarded as a propitious opening for the southern
season. In other years the season develops more gradually from tlie south,
gradually extending northwards, and being without any northern reinforcc'-
ment nunains enfeebled : in such years the inland dry regions usually experi-
ence low’ precipitations. Tlie season retires as it begins, the rainfall' systcnns
gradually weakening, until in October the precipitations affect th<'' lower
south-west only, and there is a return to drought in November.

XXXll.

Charles Austin Gardner.

The third climatic region, the vast central area, receives nothing of the
pluvial wealth of the northern ami southern systems. If, as sometimes hap-
pens, the southern system sets in over a wide front in conjunction with an
active low-pressure northern system, then the central area may receive heavy
and useful precipitations, and in such years the Wiluna district may receive
more annual rainfall than many coastal stations. On the other hand, the ac-
tive hurricanes of the months of February and March may bring widespread
rains to the north-west and extend far inland, bringing disastrous floods.
There are thus two months in the year which receive more rainfall than any
others- — March and June, the foriner being summer rain, the latter winter
rain. Having no rainfall system peculiar to itself, this central region relies
entirely upon extensions of the northern and southern systems, experiencing
sometimes a comparatively rich rainfall in March, or in June, with lighter
precipitations at other times, w'hile on the other hand, one or both of these
may fail, and a prolonged period of drought ensues, sometimes extending
over two or more years. On the other hand, it may experience two heavy
rainfall systems within the one year, as was the case at Wiluna in 1900 and
1942. It is this climatic instability that determines the character of the vege-
tation of the middle zone. The following table illustrates the annual rainfall
for two selected stations : Wiluna being typical for the middle of the dry

area, and Katanning in the South-West being typical of the winter rainfall area.

These tables are interesting in that they indicate the high fluctuations
in the annual rainfall for Wiluna when compared with the relatively reliable
rainfall of Katamiing. At Wiluna we have a departure from the moan of
-f- 201 per cent, and — ■ 79 per cent., whereas at Katanning similar figures over
the same period are H- 45 per cent, and — 26 per cent.

ANNUAL RAINFALL AT WILUNA, 1899-1942, SHOWING ANNUAL VARIATION.

(Annual average

rainfall.

00

'.O

1

931 points.

Year.

Annual

Total.

De[:>arture
from Mean.

Year.

Annual

Total.

D(' part lire
from Mean.

1899 ....

855

—

8

1922 ....

691

—

26

1900 ....

2,803

_L

(

201

1923 ....

742

—

20

1901 ...

875

—

18

1924 ....

506

—

46

1902 ....

1,366

-L

47

1925 ....

K430

-f

54

1903 ....

737

—

21

1926 ....

633

—

29

1904 ....

1,077

16

1927 ....

1.034

1

~r

11

1905 ...

617

—

34

1928 ....

460

—

50

1906 ....

780

—

16

1929 ....

805

—

13

1907 ....

804

—

24

1930 ....

1,564

-h

68

1908 ....

1,019

+

9

1931 ....

1,374

49

1909 ....

1,334

+

43

1932 ....

654

—

29

1910 ...

192

—

79

1933 ....

1,032

11

1911 ...

676

—

27

1934 ....

1,412

+

52

1912 ....

745

—

20

1935 ...

363

—

62

1913 ...

1,396

+

50

1936 ...

271

—

71

1914 ...

790

—

15

1937 ....

804

—

14

1915 ....

1,693

-L

82

1938 ....

812

—

12

1916 ....

1,061

14

1939 ....

778

—

16

1917 ....

1,086

17

1940 ....

346

—

62

1918 ....

987

+

(i

1941 ...

1,100

+

IS

1919 ....

935

0

1942 ....

2,770

1

147

1920 ....

847

—

4

1921 ....

729

—

22

The Vegetation of Western Australia.

XXXlll.

RAINFALL FOR KATANNING, 1891-1941.

(Average 1,873 points.-)

Year.

Annual

Total.

Departure
from Mean.

Year.

Annual

Total.

Departure
from Mean.

1891 ....

1,236

points
-- 637

0/

/o

34

1917 ....

2,566

points
+ 693

9/

/o

37

1892 ....

1,568

—

305

16

1918 ....

2,316

+

443

24

1893 ....

1,621

—

252

13

1919 ....

1,700

—

173

9

1894 ....

1,293

—

580

31

1920 ....

1,975

102

5

1895 ....

1921 ....

2,080

+

207

11

1896 ....

1,440

—

443

24

1922 ....

1,605

• —

268

14

1897 ....

1,274

- — ■

599

32

1923 ....

2,394

521

28

1898 ....

1,770

— .

103

6

1924 ....

1,715

— .

168

9

1899 ....

1,784

■ —

89

5

1925 ....

1,972

99

5

1900 ....

2,133

260

14

1926 ....

2,461

588

31

1901 ....

1,649

—

224

12

1927 ....

1,776

—

97

5

1902 ....

1,429

—

444

24

1928 ....

1,862

—

n

1

1903 ...

2,431

538

30

1929 ...

2,024

+

151

8

1904 ....

2,023

150

8

1930 ...

1,943

+

70

4

1905 ...

2,453

580

31

1931 ....

1,915

+

42

2

1906 ....

1,936

63

3

1932 ....

2,278

+

405

22

1907 ....

1,948

75

4

1933 ....

1,972

99

5

1908 ..

1,454

—

419

23

1934 ....

2,068

+

195

10

1909 ....

2,066

+

193

10

1935 ....

1,700

+

173

9

1910 ....

2,013

4-

140

7

1936 ....

1,525

—

348

19

1911 ....

1,610

—

263

14

1937 ....

2,215

342

19

1912 ...

1,583

—

290

15

1938 ...

1,384

—

489

26

1913 ...

2,468

-f-

595

32

1939 ....

2,718

00

Or

45

1914 ....

1,593

—

280

15

1940 ....

1,072

—

801

43

1915 ....

2,189

316

17

1941 ....

1,857

—

16

1

1916 ...

1,588

—

285

15

Mean var.— 16

•1

2. TEMPERATURE.

The annual mean temperatures show little correlation witli the vegetation
in general, but are of importance in limiting the megathermic elements. Tlie
mean isotherm of 80° F. for January and February — the driest months in tlie
soutli — closely approximates to the southern boundary of the Mulga forma-
tion, but this may be little more than a coincidence. What is of more im-
portance is the temperatin-e range, especially the diurnal range. While tlie
coastal temperatures remain fairly equable, high diurnal variations an? ex-
})erienced in the interior, especially in the arid areas ; even in the soutli-west
tliey r(‘main extreme at such stations as Bridgetown and Katanning. The
\ariation between the monthly maxima and the montlily minima are pro-
Ijortionately high, being 45° F. at Katanning, 53° at Kellerbcn-rin, and rising
to ()0° at Muiidiwindi. I’liese temperature extremes, especially the diurnal
extremes, are comparable in tlie cases of Wiluna and Mundiwindi, with those
of the desert of the Punjab ! A study of the monthly isotherms shows an
interesting correlation between the temperature and the vegetation, ami
wlien applied to the months of maximum precipitation explains satisfactorily
the extent of the grassland and woodland formations.

X.NXIV.

ChARLKS AuyTTX (tARDNER,

3. HUMIDITY, EVAPORATION, AXD WIND.

There are but 36 stations in Western Australia from which data are
l>ublished concerning temperature and humidity, and these are (jviite in-
adequate for tlie purpose of constructing a map. The appended tables have
been compiled for reference and the forming of general conclusions, and contain
all the information which has been officially published.

While the 9*0 a.m. readings for re^lative humidity may be regarded as.
typical for the 24:-hour period in certain coastal areas, it \\'ill be at once aj)-
parent that this is not so for the inland stations ; the relative humidity tables
of monthly mean percentages illustrate this point quite clearly. They exhibit
the fact that in the wet forest areas, the range between morning and afternoon
(9*0 a.m. and 3-0 p.m.) readings is very slight, while in the sclerophyllous.
woodland formations it is high, e.tj., while at Cape Leeuwin this figure does
not exceed 7 per cent., in inland localities it is as high as 24 j)(‘r cent..
Another point that has to be considered is that the 9*0 a.m. leadings taken
in summer, when in the south the sun rises at 4-30 a.m., are quite a different
(juantity from those taken in winter, when the sun rises at 7-0 a.m. It is-
interesting to compare the relative humidity figures for Kellerberrin and
Perth with those of Carnamah and Mundiwindi, all of v4iich have different
climates. The table of relative liumidity for the four consecutive driest
months is also instructive, and emjdiasises still more the climatic extremes,
of the inland stations.

Information regarding ovafjoration can be obtained from the figures given
for the saturation deficit and the Meyer Ratio. There are only four official
evaporimeters in Western Australia, the most inland of which is at Merredin.

One at Wiluna or Mundiwindi would be of much value. The following tabl&

gives the monthly evaporation

in inches

for the four

stations

-

Perth.

Narrogin.

Chapman.

Merredin.

January

10-370

9-083

1 1 - 940

13-151

February

8-650

7-570

10-192

1 0 • 860

March

7-550

6-433

9-226

9-359

April

4 • 670

3-780

6-792

5-979

May

4-670

2-361

4-160

3-641

June ....

1-770

1 -609

2-629

2-289

July

1 -730

1 624

2-473

2 - 09 a

August

2-360

1 -864

2-942

2-611

September ....

3-410

2-513

4-055

4 - 010 '

October

5-340

3-833

6-431

6-527

November ....

7 ■ 660

6-156

8-951

9-764

December

9-780

8-505

11 -391

12-285

Total

65-070

55-331

81-162

82-557

4. RATIO OF PRECIPITATION TO EVAPORATION.

AVliile heat and moisture are the most important of the elements of the
climate affecting plant life, water is of importance to the plant in tvs o ways :
as soil water, and as atmospheric moisture. The relative humidity depends,
upon the temperature of the air and the water vapour present ; the hotter
the air, the more water vapour it can hold, and the relation of the temperature
of the air to its water vapour content controls the loss of water from the aerial
j)arts of the plant. Vegetation is therefore dependent upon precij^itation
and evaporation.

The Vegetation op Western Australia.

XXXV.

Tlie evaporating power of the air is expressed by what is termed the
^saturation deficit. This is the amount by which tiie partial pressure of water
vapour in a given volume of air falls short of the pressure at saturation point,
and is thus a direct measure of tlie eva}Dorating power of the atmosphere.
Tn the appended tables, the mean annual saturation deficit values are given,
•as well as figures for the four consecutive wettest months. They are useful
for })urposes of comparison, and attention should be paid to the relative
values for inland stations.

The Meyer Ratio, which is the ratio of precipitation to the saturation
<deficit (expressed as an annual ratio), shows a high dc'gree of correlation with
Precij)itation /P\

- 7 T 1 - I, and has been omploved in sevwal countries as an ecological

Evaporation \E/

index, proving of much value in this respect. Mapping the distribution of
this ratio with any degree of accuracy is almost impossible, since there are
only 3f> stations in Western Australia from vhich data are }>nblished. The
accompanying map may, however, prove of interest, although the absence
of recording stations in the Fortescue district is unfortunate. What the
map does show, however, is that the Meyer Ratio determines the approximate
boundaries of the forest formations of the south-west, and there is a general
correlation between its isoj:)leths and the boundaries of the vegetation pro-
vinces. In the north, the 50 isopletli may be taken as representing the south-
cun boundary of the monsoon woodland area.

The

Seasonal precipitation / SP

VSSD/

Ratio is also given for

com-

Seasonal saturation deficit
parison with the Meyer Ratio, with which it shows a certain correlation, but
serves to emphasise still further the thermo-illuvial factor. This ratio is the
seasonal equivalent of the Meyer Ratio, and as such perhaps serves as a better
climatic index in areas which are characterised by a seasonal period of drought.
Seasonal Precipitation /SP'

The

/o.r\

I Ratio is also important in indicating

Seasonal Temperature
types of vegetation, but is mainly of value only when considered in conjunc-
tion with some other factor.

Finally, amongst the climatic statistics, nn important consideration is
tlie variability of the annual rainfall. In the accompanying tables this is
c'xpressed as a percentage over the number of years during which records have
been taken. It serves as an intlex for the drought factor, and in this respect
is important when considering the climates of the Northern and Southern
Provinces in contrast to the Eremea.

Included in the climatic factor, atmospheric movements are of con-
siderable importance. Especially in the dry interior, the value of wind as a
desiccating agent determining the habit and structure of plants is very con-
siderable. Any casual study of the vegetation will at once illustrate this
fact. These winds are mainly of two types : tlie dry prevailing winds that

dominate almost the entire interior, and the strong sea winds that mould the
structure of certain coastal formations. These latter appear to exert their
most powerful effects on the trees of the littoral between Oeraklton and Don-
garra, and on the dune thickets of the south coast. Their influence is more
fully discussed in the cha[)ter dealing with growth-forms.

VI.-

RELATIONSHIP8 BETWEEN THE C^LIMATE
OF WESTERN AUSTRALIA.

AND VEGETATION

From what has already been stated it will be evident that Western Aus-
tralia can conveniently be dividetl into three climatic provinces determined

XXXVI.

Charles Austin Gardner.

by the seasonal incidence and amount of the rainfall. The thermic factor is also
very important in distinguishing the northern from the southern rainfall
areas, while the middle area — the Eremea— stands as a buffer between the other
two.

The main purpose of this thesis has been to define as clearly and as
accurately as possible, the three vegetation provinces and to correlate them
with the climatic provinces, for, of all the factors which determine and control
the vegetation of the earth, the climatic factor is the most important, and
rainfall and temperature are the most important of the climatic elements.

The natural vegetation of a country, especially when unaffected by the
biotic, or any other external factor, is the index of its climatic and edaphic
factors. In Western Australia, where, in the virgin state each association
represents an ecological climax, it is possible to learn much concerning both
soil and climate from the study of the vegetation. In fact, it is possible to
obtain a more or less clear picture of the climatic conditions by such a study.
Further, there is a close correlation between the soil formations and the climatic
types, and soil surveys in Australia are to a great extent based on the vegeta-
tion.

In the Introduction we discussed briefly the elements of the vegetation
of Western Australia, showing that in the north, under conditions of summer
rainfall and a prolonged season of drought, a type of \*egetation existed in
which the Palaeotropic Element was at home. In the south-west, an area
described as tlie “ triangular-crescentic area extending from Shark Bay to
Israelite Bay ” came under the influence of the autochthonous element,
largely made up of the Antarctic and Australian Elements. It is the region
which comes under the active influence of the winter rains, with a summer
period of drought. The third area, which we call the Eremean Province, is
an intermediate area characterised by a capricious climate, of uncertain
regularity, of extreme temperatures, and dependent upon the rainfall systems
of the neighbouring provinces for what rain it receives. This area we found
to be dominated by the Neo-Australian Element, which is derived from botli
southern and northern sources of origin.

The boundary of the South-West Province can be determined within
comparatively narrow limits, allowing for certain edaphic variations, and is
primarily determined by the gradual diminution of the winter rainfall, partly
by teniperature, and to a great extent by the seasonal variability of the rain-
fall.

As we pass from the coast towards the interior we notice a cliange in
the formations, from high forest, through v'oodland and heath to mulga-
bush and ultimately steppe and desert. Ihit quite apart from these changes,
there is a graduation in the floristic composition. As we approach the bound-
ary of the South-West Province we notice that the autochthonous species
become gradually more rare, first we are aware that such plants as Anigo-
zanthos and Conostylis gradually disappear, and in turn the Epacridaceae
cease. Gradually we find an increase in the species of Eremophila, of certain
Chenopodiaceae, and in the woodland the Zygophyllaceae and Compositae
become increasingly important, together with a more open formation of the
woodland ground flora. The actual line of the boundary can be determined
by the ephedra-like species of Eremophila, Zygophyllum, and Chenopodiaceae,
while in the heath formations we find the last of the south-west elements
in Boronia and Epacridaceae. These, and some other indicative elements
announce that we are passing from the South-West to the Eremea. Quite
apart from changes in the soil, this is brought about by the gradual diminution

The Vegetatiox op Western Australia.

xxxvii.

in the rainfall, especially the seasonal rainfall. The following table illus-
trates this diminution : —

Station.

Altitude.

Distance E.

or W.

Rainfall

ft.

of escarpment.

(points).

Fremantle

C

20

miles

W.

3,027

Perth . .

197

10

w.

3,467

Guildford

30

5

> i

w.

3,393

Mundaring

975

6

if

E.

4,352

Chidlow

983

10

i!

E.

3,796

Northam

494

40

ii

E.

1,711

Meckering

640

60

a

E.

1,526

Cunderdin

733

75

a

E.

1,458

Tammin

800

85

a

E.

1,378

Kellerberrin

815

100

ii

E.

1,329

Doodlakine

840

110

a

E.

1,235

Merredin

1,042

135

ii

E.

1,292

Walgoolan

1,055

155

ii

E.

1,196

Southern Cross

1,157

200

i i

E.

1,052

Coolgardie

1,394

315

ii

E.

1,009

Kalgoorlie

1,234

337

ii

E.

951

Rawlinna

603

570

ii

E.

637

Loongana

603

680

ii

E.

615

A corresponding diminution occurs everj"N\’here ; to the east of Geraldton,
for example, and to the north of Esperance, the diminution takes place to a
greater degree over a given area, and the example quoted above is the longest
transect possible with so many stations. It is this gradual diminution that
•accounts everywhere for the gradual changes in the vegetation, modified only
by the edaphic factor.

In constructing a climatic map of Western Australia which would con-
form to the main types of vegetation, it was obvious that any consideration
of the annual isohyets alone would be insufficient. For amongst other con-
siderations, such rainfall statistics do not take into account the useful seasonal
rainfall, and give no indication of seasonal periodicity, nor do they indicate
those areas which experience a pronounced drought factor in the annual
climatic cycle. In other words the essential elements of temperature and
rainfall periodicity are not indicated, althougli in a very general way there
is some correlation between the annual rainfall figures and the principal types
•of vegetation. What is required therefore is a map embodying both pluvial
an<l thermal factors, but this is impossible, because the temperature figures
wei*e obtainable from 30 stations only.

A means of employing the temperature factor was reached by considering
the seasonal incidence of the wettest period of tlie year. At first the six
M'ettest months were employed, but were later abandoned in favour of a season
employing the four wettest consecutive months, and the four consecutive
<hiest months. The isohyets obtained by a consideration of the four wet
months, referred to throughout the tables as the '"season,” were fo\md to co-
incide very closely with the boundaries of the three provinces of the vegetation.
Tlie line of 175 mm. winter (May-August) rainfall is, in fact, exactly a very
distinct line of separation, cleai’ly defining the limits of the autochthonous
elements of the south-west, and also determining the western limits of those
elements proper to the Eremea. It is tlie Western Australian equivalent of
the “ Goydei' Line ” of South Australia. Apart from its significance in the
vegetation of Western Australia, it may well prove to be a line of consideraVile
importance in future agricultural development.

XXXVlll.

Charles Austix Gardner.

Tlie same \'alue in the seasonal isoliyets of the northern wet season (the
four conseciitive months which lie between December and April, 7.e., December-
Mareh. or January -April), 175 mm., was found to correspond very closely with
Avhat is considered to be the boundary of the northern sav'^annah types, and aLso
to conform to the limits of many Indo-Malayan groups in northern Australia.

The third climatic province is that in which the rainfall for the four con-
secutive wettest months is less than 175 mm. It will be noticed here that
these four months are not uniform over the area, but that the winter system
(May-August) extends along the southern and western margin of the Province,
as the December-March or January-April season does in the north. The
middle zone howev'cr, experiences its four wettest months between March
and June, exactly those months which experience the northern and southern
maximum extensions of precipitation. The intervening months, April and
May, have a lownr rainfall than either March (summer or northern) or June
(southern or winter) rainfall. This seasonal incidence has, of course, a power-
ful thermic factor which is reflected in the relative importance or absence of
grass in the formations. Whereas the Northern Province is essentially the
area in \vhich the Indo-Malayan Element is important, and grass the pre-
dominant ground flora, and the Soiith-West Province the area deficient in
grass, but in which the autochthonoxis flora holds undisputed sway, the
characteristics of the intermediate Eremean Province correspond to an area in
which the flora shows affinities to the regions both to the north and south,,
admitting both to a certain extent, but liaving at the same time its own dis-
tinctive impress, forcibly expressing the capriciousness of its climatic factor
and preseiAung a strange duality in its herbaceous species- — -a tendency to-
wards grass under warm influences, and dicotyledonous herbage under cooler
conditions of moisture.

The three climatic Provinces thus delineated are : —

The Northern Province, the South-West Pro^dnce, and the Eremean
Province.

VII. THE PPOVINCES AND THEIR FORMATIONS.

1. THE NORTHERN PROVINCE.

The Northern Province is cliaracterised by the role whicji is played by the
Indo-Melanesian Element, by its formations, in vhich grass takes a prominent
part as the most important ground flora, and finally by the predominanc(^
of megathermic plants. Except in the riverain and mangrove formations^
grass occurs almost everyv'here. Sclerophyllous shrubs are in the main either
entirely absent or very scarce, and the trees are usually sparsely distributed.
In tlie southern dry areas Triodia steppe and Pindan occur, with savannah
on the alluvial soils. Only where laterite is present do we find any attempt at
a sclerophyllous formation, and these formations bear a distinctive physiognomy
by reason of tlie frecpient occurrence of palms and broad-leaved shrubs. Even
here the coarse grasses assume an importance which causes a sharp contrast
with similar formations in temperate latitudes.

It is interesting to consider briefly the role taken by the Indo-Melanesian
plants in this Province. Apart from the mangrove formations which fringe
the coast and become of sufficient size and density to constitute a forest in the
estuaries, the real home of this Element is to be found in the riverain forests-
where, under the influence of telluric water, species which are more properly
those of the rain forest find a suitable environment independent of atmospheric
I^recipitations. Tliis is especially true of the black muddy soils, or soils rich in
humus. \Mu‘re the river flats are well drained, the formation ceases to exist.

The Vegetation’ of Western Austk-alia.

XXXIX.

The occurrence of epiphytic orchids, tall climbing ferns, dense forests of cad-
juput (Melaleuca leucadendron) and many broad leaved trees such as Sarco-
cephalus, Ficus, Randia, Seshania grandiflora, Barringtoyiia, and Carallia,
with occasional species of broad leaved Terminalia, renders the formation at
once distinctive, especially when the density of the canopy is considered and
the predominance of species of Pandanus and ground flora of ferns.

The monsoon woodland occurs in the higher rainfall regions, but is most
highly developed in the sandstone areas. Apart from the Eucaly2:)tus species
which form the greater part of the arborescent growth, there is a distinctive
IndO'Malayan impress by reason of the number and density of Indo-Malayan
arborescent plants, such as Terminalia, a number of Apocynaceae and Rubia-
ceae, e.g., Timonius ; species of Diospyros, Maha humilis, Planchonella, and
Lucuma, and many smaller shrubs, with Cycas and groves of Livistona. Im-
portant amongst the species of Eucalyptus are E. ndniata, E. tetrodonta, E.
terminalis, and E. hrachyandra. Deciduous species of Sterculia and Brachy-
chiton are also common. In fact, the deciduous character of the foliage of
many species, including Eucalyptus hrachyayidra , gives to the formation an ap-
pearance which is unique amongst the formations in Western Australia,

The savannah woodland of Kimberley, which extends generally southwards
to the Fitzroy River, has much in common with the monsoon woodland, but
is much more open in composition, with a greater abundance of perennial
grass, and Eucalyptus plays a more important role. There arc two principal
types : — the basaltic savannah, and the sandstone savannah. The former
consists of almost pure stands of Eucalyptus Spenceria,na , associated with
Eucalyptus clavigera, and the less important scatterc'd trees of Eucalyptus
latifolia and a few minor species. Gossampinus hepiaphylla, Erythrina ves-
pertilio, and species of Terminalia are amongst the most important deciduous
species, together with Sterculia. There are \’(‘ry few shrubs, with the ex-
ception of Goclilospermum and Maha, and the ground is densely covered witli
perennial grasses, notably Themeda triandra. The trees are well spaced, and
apart from the species of Eucalyptus, the only prevalent evergreen trees
are E^'ythrophloeum., Hakea, and Grevillea. The Indo-Malayan Element is not
well represented, except in the herbaceous species, and floristically it is the poor-
est of all the tropical formations.

The sandstone savannah, on the other hand, is much richer floristically.
It does not possess the uniform character of the basaltic savannah, and there
is a more marked intrusion of the Indo-Malayan Elemtmt. E‘ucalyj)fus termhi-
aha, E. tetrodonta, and E. miniata are amongst the commoia^st of the Eucalyp-
tus trees, with isolated groups of Eucalyptus alba' on the low-lying flats. The
Proteaceae are represented by Persoonia falcata. sevei-al specaes of Grevillea
and Hakea, with Grevillea chrysodendron and Bmtksia dentata in the swampy
depressions. Rut it is above all the strictly tropical groups that give to the
sandstone savannah its distinctive character ; Celasti'us, Denhamia, Ficus,.
Terminalia, Etigenia, Careya, Celtis, Maha, and many other woody plants,
including the common Carissa lanceolata b('ing met with almost everywhere,
together with the deciduous Sterculiae and the grotes(jue Baobab tree (Adan-
sonia, Gregorii), Th(^ herbaceous and sub-slirubby plants are represented here
to a degree not found elsewhere in the tropics, the bidliant flowers of Hibiscus,
Cienfugosia, and Thespesia being amongst the most conspicuous of the Mal-
vaceae and Tiliaceae, which are predominant. The herbaw'ous flora, too,
IS rich in widely diversified species and growth forms. Mfliiie the commonest
plants are tlie gi’asses which occur in splendid profusion, the Asiatic, Aus-
tralian, and African groups being mixed together, but tisually pr('dominat('d

xL

Charlkr Austin Gardner.

by species of Sorghuin, one of which (S, intrans) attains a height of six or
eiglit feet. There are also a multitude of dicotyledonous herbs, including
Ptilotus and Trichmum, Spermacoce, many species of the Scrophulariaceae,
Leguminosae, and plants belong to the Centrospermae.

The swampy areas are mainly covered by Cyperaceae belonging to many
genera, wdth Mitrasacme , Byhlis, and Utricularia, w'here the soil is perennially
moist.

Whereas the basaltic savannah woodland maintains its distinctive physi-
ognomy throughout, tlie sandstone savannah woodland, on account of the
deeply eroded nature of the country^ is remarkably diversified. No large
uniform areas are to be seen. Either the plateau is flat, or it is seamed with
wild deep canyons, and wide eroded flats. The swampy land referred to occurs
usually in depressions in the plateau itself, or in the broad valleys flanked by
■steep clifls. The canyons and gorges carry a profusion of lithophilous shrubs
or trees, such as Ficus, Erioglossum, Dodonaea, Boronia, Calythrix, and the giant
Verticordia, Ervatamia, Solarium, and a number of species in which the tem-
perate Australian groups are mixed with the tropical. The liigh plateau, on
the other hand, supports plants w’hich are more typical of the dry steppe
zone~Triodia, Grevillea, Callitris, Dampiera, Cassia Eucalyptus papuana,
-and E. setosa, and Acacia species of the Phyllodineae, notably A. tumida,
A. Luehmanni, and A. sericata.

This contrast is remarkable, and exhibits the fact that the Indo-Malayan
Element in the flora finds its highest expression in the low-lying moist soils,
while the more arid situations are almost entirely populated by xerophytic
Australian species which are alone able to thrive in this unfavourable environ-
ment, for it must be remembered that although the Hann Plateau of Kim-
berley receives a rich rainfall in its wet season, the six cool months experience
a drought period scarcely equalled, even in the Eremea, and that the meso-
phytic and hydrophytic elements of the flora, including these Indo-Malayan
species that are not deciduous, are restricted to soils in which telluric water
supplies their moisture requirements.

The King Leopold Range marks the southern boundary of the savannah
woodland, and we enter into a region of lower rainfall. The principal formation
is savannah carrying isolated or scattered trees, mostly Eucalyptus and Acacia,
■and a dense ground-covering of grass and herbs. Where the soil is rich in qual-
ity, or subjected to periodic inundation we find a richer flora, in which a number
of tropical African grasses grow in abundance mixed with genera which are
either gerontogeous, or with a few strictly Australian t^’pes, together with sev-
eral Indo-Malayan or paleotropic herbaceous species, especially legumes. Here
we also find bipinnate Acacias, especially A. Farnesiana and A. Bidwilli, and
the outposts of the Indo-Malayan arboreal Element — plants such as Bauhinia,
Pithecolohiimi, Dichrostachys, and a few others of less importance. The savan-
nah is only well represented on the plains, especially in alluvial soils. Where
the soils become more sandy, or of a gravelly nature, the physiognomy changes,
and we enter into the Pindan country near the Fitzroy River. The Pindan
country is perhaps the tropical equivalent of the Mulga bush of the Eremea,
but its woody plants, mostly Acacia, are more densely crowded, and the Euca-
lyptus trees become more depauperate. E. setosa and other Corymbosae are
the dominant species. Associated with the shrubs and dwarf trees are tall
coarse grasses and succulent lierbage, such as Portidaca, Trianthema, and Sesuv-
ium. In its more impoverished form it leads to the steppe and ultimately
to the desert some distance to the south of the Fitzroy River.

The Vegetation of Western Australia.

xli.

The main area of the Northern Province is connected with the Fortescue-
De Grey region by a narrow neck of Pindan country in the vicinity of Wollal.
The Fortescue-De Grey region, which centres round the Hamersley Range
has been referred to previously as an area which is almost an “ island ” in
the Eremea. The whole of this area is not well known phytogeographically,
but the extensive development of savannah, the climatic conditions, and the
existence of many vestiges of the Indo-Malayan Element, render its inclusion
within the Province necessary. There are, however, several important
changes. The deciduous species no longer exist, with the single exception
of Brachychiton australe, but the prevalence of such plants as Owenia, Livistona^
Atalaya, Bauhinia, Dichrostachys, and a number of others, including certain
lithophytes and mesophytic herbs, shows that this country has many re-
semblances to that of the north. In the savannah. Sorghum, Eulalia, Chryso-
pogon, Heteropogon, Iseilema, and Astrebla remain as important constituents —
plants which are almost totally absent from the Eremea. Here again the savan-
nah is restricted to the alluvial soils, merging into a type of Pindan and the
Triodia steppe. Along the rivers and in the declivities of the Hamersley Range
we find such plants as Astrotricha, Ficus, Terminalia, and Clerodendron, whilst
the Rubiaceae and Scrophulariaceae, together with several megathermic
Compositae, remain in sufficient numbers to be regarded as important. The
actual western boundary of the Northern Province near the Ashburton River
is not known, but occurs near Mardie m the west, and includes Red Hill, while
to the east it extends to Marble Bar and Nullagine. Whether or not the
Province extends to the south of the Hamersley Range is not known with
certainty.

2. THE SOUTH-WEST PROVINCE.

The South-West Piovince is distinguished floristically by the rich develop-
ment of the Australian and Antarctic Elements. This c'e/elopment is most
powerfully expressed towards the cusps of its tringular-crescentic area of
distribution, both in the north, between the Hill River and the Murchison
River, and in the south-east between the Phillips River and Israelite Bay.
There is, indeed, a close connection between the plants of these two widely
separated areas, as for example the restricted distribution of Phymatocarpus,
Conothamnus, and certain species of Eucalyptus and Adenanthos, as well as
certain close affinities in a few groups such as Conostylis and Anigozanthos.
Both areas represent the richest areas in species and forms in the entire pro-
vince. It is also noteworthy that these two areas are precisely the regions
where the climatic and epharmonic gradients are the sharpest, and thus com-
petition between Eremean and southern forms least possible of extension.

Other characteristics of the South-West Province are the predoininence
of sclerophyllous shrubs in every formation, either as undergrowth in the
forest and woodland, or crowding the heaths. The floristic wealth of these
heaths is probably unparallelled an;ywrhere else on the earth. Important,
too, is the paucity of herbaceous plants, particularly the grasses, although
the Cyperaceae, Restionaceae, Haemodoraceae, Liliaceae, and Compositae
make up to some extent for this deficiency. When one considers the facility
with which certain exotic grasses establish themselves, as for example in
the tuart country, this paucity of the indigenous grasses is difficult to under-
stand. It may be the story of the warfare which is supposed to exist between
woodland and grassland, and that the climatic conditions favour the develop-
ment of trees and shrubs, or it may be a question of the biotic factor, or of
species migration, for the greater number of the indigenous grasses, with
the exception of a few such as Festuca, Danthonia, Stipa, and Amphipogon^
appear to be of palaeotropic derivation, and have not been able to obtain a foot-

xlii.

Ctiaiu.ks Austin Gardxkr.

hold in the South-Wt'st because of their inability to surmount the climatic
barriers. In any case, tlie number of genera of grasses in the South-West
Province is exceedingly small, and the area possesses but one endemic genus,
the monotypic Diplopogov .

Lastly, it is the liigli degree of endemism within the Province that is one
of its outstanding characteristics. Tliis has already been remarked upon, and
the figure is about 80 p('r cent. L'^nlike the Northern and Eremean Provinces
it has borrowed little or nothing except from the ancient ancestors from which
it has evolved, and of course excepting those maritime or littoral plants whicli
have suffered oceanic transportation, and which have obtained a foothold in
an (mvironment whicli is not typical of the general area.

Apart from the littoral halophytic and dune formations which require
no particular attention beyond tlie fact they they contain more or less cos-
mopolitan maritime, or Eremean elements, the south-western formations are
mainly woodland and lieath types. In the extreme south-w'est, in the area
bounded by the 500 mm. seasonal isohyet, the forest formations predominate
except where sandy coastal plain or swamps provide unsuitable edaphic
conditions. To the east of this area, between the seasonal isohyets of 500 mm.
and 200 mm., the savannah woodland and heath formations predominate, while
below the 200 mm. isohyet, sclerophyllous woodland and sand heath dominate
the landscape. While the above isohyets may be said to determine broadly
the limits of these formations, the edaphic factor is important in determining
the formations under drier conditions. For example, while the edaphic factor
is not so active in the forest areas, it does to a considerable extent determine
the sclerophyllous woodland, savannah woodland, and heath formations, the
sclerophyllous woodland always being determined by the soil, thus dove-
tailing into the climatic confines.

Formations which are more strictly edaphic are those of the swamp, the
laterite, and the granite. It would, however, be cpiite beyond the scope of
this essay to deal with tliese at any length, and the merest outlines must
suffice.

(a) THE FOREST FORMATIONS.

(i) The Jarrah Forest.

The distribution of Eucalyptus marginata is bounded by the 400 mm.
seasonal isohyet, but it is only on the lateritic soils that the forest formation
exists, for although it occuis freely on the sandy soil of the coastal plain, it
does not in this envii'onment constitute a forest. The jarrah forest is a true
sclerophyllous formation, remaikable for the paucity of other tree species, for
it is not a social species, and its only associates are Eucalyptus patens and E.
calophylla. At the same time, this is surely one of the outstanding character-
istics of the Eucalyptus associations. These trees are rather widely branched,
but suffer little oi‘ no ovei'lapping of their crowns, the result being a forest
canopy easily penetrated by light. Amongst the Eucalyptus trees are smaller
trees, mostly Myrtaceae. Proteaceae, and Casuarinaceae, e.g., Persoania,
Eanhsia. Xylomelum^ Casuariua, and Agonis, the only one of which that
exhibits any marked degree of plagiotropism being Banksia grandis, and to
a lesser extent Xylomeliim occidentale. The undergrowth is composed of
sclerophyllous slirubs from less than a metre to two or three metres high,
pi’edominantly Mvrtaceae, Protraceae, and Leguminosae. In certain areas
Macrozamia becomes impoitant, together with Xanthorrhoea^ while towards
the south the remarkable suffrutescent Podocarpiis Drouyyiiana assumes im-
portance.

The Vegetation of Western Australia.

xliii.

Floristically, however, the formation is not richly endowed. The herb-
aceous plants are poorly represented, and there are but a few lianes, mostly
Hardenbergia, Kennedya, and Clematis. It has been stated previously that
the lateritic formations are usually poor in species, and this rule holds good
for the jarrah forest. The forest attains its best development in the vicinity
of tiie Blackwood River, where the undergrowth, due to a heavier canopy,
becomes less dense, and Pteridium becomes important in the lighter soils.
The jarrah and its competitive trees are rough barked, a (iharacteristic which
is unusual amongst the trees of southern Western Australia.

(ii) The Karri Forest.

Tlie karri forest stands in sharp contrast to the jarrali forest by reason
of its larger trees, the mesophytic undergrowth, and the better development
of an under-storey of trees, in which Casuarina decussata, Agonis flexuosa,
Banksia, and Hakea take a prominent place. The shrubby undergrowth is
higher, Acacia pentadenia one of the Bipinnatae, with large-leaved shrubs
such as Trymalium, Chorliaena, Hovea elliptica, and Albizzia, and in the open
spaces, areas of tall Pteridium, often exceeding two or three metres in height.
The sclerophyllous plants of the jarrah forest have larger, less sclerophyllous
leaves, and the smaller shrubs and herbs are of straggling widely-branched
habit. Indeed, the wide branching and interlocking of branches is at once in
marked contrast to the shrubs of other formations, as well as the tendency
for species to become ombrophytes. The principal shade is, however, not
<lue to the density of the canopy provided by the principal trees, for although
these, the karri {Eucalyptus diversicolor), tingle trees {E. Jacksoni and E.
Guilfoylei), and Marri {Eucalyptus calophylla) are precisely those that have
the widest and most horizontal branching habit of all the south-western species
of the genus, they do not give much shade. The shade is provided by the
under-storey of smaller trees, especially Casuarina decussata and Banksia
grandis, or the other species of Banksia. The true ombrophytes are those
species w'hich live on the forest floor, such as Dampiera hederacea, the species
of Boronia, Tremandra, and Opercularia.

Unlike the jarrah forest, the karri forest secures its edaphic requirement
in the soils derived from the granitoid or gneissic rocks which more or less
determine its occurrence in its general area of distribution, and account for
outlying areas on the Porongorup Range, and near Mount Manypeak. The
karri forest occurs bet-ween Denmark and the Blackwood River estuary,
where the rainfall during the six driest months exceeds 200 mm. The density
of the vegetation, both amongst the arborescent or fruticose species, together
with the occasional occurrences of epiphytic ferns, gives to the karri forest a
physiognomy not possessed by any other plant formation in Western Aus-
tralia except the riverain forest of the Kimberley Division. It is really a type
of temperate rain forest.

(iii) The Tuort Forest.

The tuart forest is a type of savannah forest in which the trees of the
tuart {Eucalyptus gomphocephala) attaining a height of 40 metres, dominate
tlie forest in almost pure stands, being only rarely associated with such
s]iecies as E. calophylla and E. cornuta. It is edapliically confined to the
littoral limestone of the western coastal plain, the range of the species ex-
tending from near the Hill River to the Sabina River near Busselton. It is
only in the southern jiart of its range, w'-here the summer (November-April)
rainfall excieeds 125 mm. that it attains to the proportions of a forest in its
species. Here it is a true savannah formation, with an understorey of

xliv.

Charles Austin Gardner.

Agonis and Banksia, Melaleuca and Hakea, with a herbaceous ground
covering in which the shrubs are comparatively few. To the north the
undergrowdh becomes shrubby, but retains something of the naturt' of
savannah, and when the shrubby undergrowth is destroyed there is a ten-
dency towards the development of grass at the expense of the shrubby growth,
thereby indicating a succession in the direction of pure savannah forest.
In contrast to the jarrah, karri, and marri, the tuart tree is of the " box ”
type, the real environment of which is the savannah formation. (“ Box ”
is a vernacular term applied to Eucalyptus trees with entirely rough grey
bark.)

(b) The Woodland Formations.

(i) Savannah (temperate) Woodland,

In the granitic soils of the jarrah forest of the Darling Range the wandoo
tree (Eucalyptus redunca var elata) occurs where the subsoil is of a clayey
nature. Here it never dominates the forest, but is found in small numbers
in the gi’anitic areas. To the east of the 385 mm. seasonal isohyet however,
it becomes the dominant tree, usually associated with Eucalyptus foecunda
var. loxophleha and Acacia acuminata, forming open savannah-like country
with a low shrubby or herbaceous undergrowth. It is usually associated
with shallow granitic soils, but extends into deeper clay soils, especially in
the south, or into sandy country to the east and north, ultimately (about
the 200 mm. seasonal isohyet) becoming associated with decomposed laterite.
Towards the northern part of its range the species is increasingly associated
with E. foecunda var. loxophleha, and Acacia acuminata, giving place to these
two latter species entirely in the latitude of Arrino, to the nortli of which
the savannah woodland formation is composed almost entirely of these two
species as far north as Northampton. To the south, in the vicinity of Wagin,
the character of the savannah woodland undergoes a change, and Eucalyptus
occidentalis becomes important, especially in the low-lying clay soil, while
the wandoo becomes restricted to the sand, until finally, in the vicinity of
Cranbrook, the wandoo disappears entirely, and its place as a savannah tree
is taken by both Eucalyptus occidentalis and E, cornuta, the former in the
clay soils, and the latter confined to granitic areas. Between Cranbrook
and Gnowangeru[) the zone limited by the 385 and 200 mm. isohyets curves
sharply eastwards, to extend along the south coast, and here the two trees
continue as far east as the Russell Range, forming in places areas of o]jen
grassy granitic country in which either or both trees occur. This is country
more suitable for grassland development than the area to the north, on
account of the higher summer precipitations. The formation ceases to exist
to the east of Israelite Bay. Throughout the whole extent of its area of
distribution it is intersected by zones of sand-heath.

(ii) The Sclerophyllous Woodland,

Somewhere in the vicinity of the 225 mm. isohyet the loamy soils support
a type of woodland which is in contrast to the savannah woodland because
of the density of its arborescent species, the greater development of shrubs,
and the paucity of grasses except those of a harsh tussocky nature. The
formation usually occurs on low-lying soils, and the woodland either consists
entirely of trees about 25 metres with an undergrowth of shrubs about a
metre higli, and bare soil beneath, or there may be a development of slimbby
Eucalyptus species known as mallees. The formation extends as far east
as Zanthus, and northwards to the mulga bush formation, but its eastward
extension is sharply limited by the limestone soil of the Nuliarbor Plain,

The Vegetation of Western Australia.

xlv.

In the western area of its distril^ution, between the 200 and 150 mm.
isohyets the arborescent species are few, comprising three principal species
— Eucalyptus salmonophloia, E, lonqicornis, and E. sahfbris, but to the east
there is a greater admixture of the species, at least a dozen trees entering
into the composition. Within the South-West Province, i.e., as far as Mer-
redin, there is a comparatively rich development of shrubs, amongst which
Acacia and Proteaceae are predominant, but to the east of the provincial
boundary there is a notable development of Myoporaceae and Chenopo-
diaceae, the former being shrubs 3-5 metres high, mostly with an ephedra-
like habit, while the Acaciae become fewer, and the woodland undergrowth
more sparse and more and more given over to Chenopodiaceous species.
Where the soil becomes more alkaline, the Chenopodiaceae become increas-
ingly important, often associated with Amarantaceae and shrubby Compositae.
Tliis is particularly the case to the east and south of Widgiemooltha, where
the grey-bush (Cratystylis conocephala) assumes great physiognomic im-
])ortance, and Melaleuca becomes more common, while the tree species be-
come more social than in any other formations, but all have more or less
the same characteristics, so that field recognition becomes difficult. For
example, at Norseman there are at least fifteen Eucalyptus species in close
association.

The sclerophyllous woodland formations are everywhere limited by the
edaphic factor — ^nowhere do they occur on sand — -and typically they inhabit
the alluvial depressions, the broad red-soil plains, or finally the alkaline soils
of the “ lake ” country. The last-mentioned type is to be found principally
in the Lake Brown-Yilgam country, southwards to Kondinin and Lake King,
while to the east, there is a broad area of the formation between Widgiemooltha
and Dowak, extending some distance westwards, and eastwards almost to
Balladonia. The soil is more or less saline, and the Eucalyptus species are
in the main different from those of the red clay soils. Eucalyptus Brockwayi
and E. leptophylla take the place of E. salmonophloia ; Eucalyptus campaspe
the place of E. salubris, and E. Flocktoniae and E. oleosa var transcontinentalis
the place of E. longicornis. Besides the low undergrowth of Chenopodiaceae,
there are larger shi'ubs, such as Melaleuca pauperifiora and M, quadrifaria,
known as Boree ”, and extensive tracts of Cratystylis conocephala, as well
as several broom-like species of Eremopkila, Where the red clay soil woodland
occurs there is a notable reduction in Chenopodiaceous plants, and the under-
growth is lower and of a more uniform stature. Acacia and Myrtaceous species
predominate.

A curious fact which is worth mentioning here is that with the increase
of summer rains to the south of the Coolgardie district, and in the Eyre district,
the grasses become of increasing importance. For example, already at Dundas,
grasses are important in the undergrowth of the open woodland, and between
Mount Holland and Dowak it is not uncommon to find, after burning, especially
with summer rains, areas of woodland country carrying a fairly dense covering
of grass. This state of things represents a succession in the woodland which
appears to be only temporary, for ultimately the Eucalyptus seedlings grow
into trees and shrubs, and the country reverts to woodland. At the same time,
areas occur which appear to be permanently clothed with grass ; these small
areas represent natural fields in the woodland, and appear to be a permanent
feature of the southern sclerophyllous woodland zone. The best known
example is perhaps Grasspatch, an area of a few hundred acres, which in its
virgin condition consisted entirely of grasses, closely surrounded by the wood-
land. Other examples of these grassy “ oases ” occur between Norseman
and Balladonia, but are mostly little more than a few acres in extent. They

xlvi.

Charles Austin Gardner.

are, however, important since they indicate that under conditions of repeated
firing or clearing, there is a distinct tendency towards a grassland succession.

(c) LATERITIC FORMATIONS.

Apart from the jarrah forest, the lateritic formations in south-western
Australia do not support arborescent species with the exception of two types.
The first is the w^andoo formation, which in the eastern area of its range on
the fringe of the savannah formation occurs on lateritic rises as a small tree
forming open woodland with a characteristic undergrowth in which certain
species of Acacia^ Callitris Morrisoni, and Casuarina campestris are character-
istic plants. This passes almost imperceptibly into a type known as “ Wodjil
country,” in which Acacia becomes predominant (especially Acacia Beau-
verdiana, A. signata, A. neurophylla, and other species of similar habit — all
rigid erectly-branched markedly sclerophyllous shrubs. Casuarina campestris
is also a characteristic species, either forming low' dense thickets, or in more
open formation associated with Acacia, HaJeea multilineata, Dodonaea, and
certain Proteaceae. This formation remains poor in herbaceous species. It
extends from the eastern margins of the South-West Province into the Eremean
Province, even to the boundary of the mulga bush.

In the w^estern part of the savannah zone, bordering on the jarrah forest
there are two important types, more or less restricted to hills and rising ground
— the isolated w^oodlands of Eucalyptus accedens, and the Mallet country.
The former enters to a certain extent into the composition of the marginal
jarrah country wdiere the subsoil is clay, and sometimes is found in association
wdth wandoo itself. It attains to its highest development betw’een Toodyay
and Williams to the w’est of the Great Southern Railw'ay, and again between
Bindoon and Mogumber. The type is characterised by its stony soil, and
the density of the sclerophyllous undergrow^th in contrast to the pure wandoo
formation.

Mallet is the name given to two or three species of EucalypUis, especially
the brown mallet {Eucalyptus astringens) and the blue mallet (E. Gardneri).
These trees, w'liich grow* in dense formation, occupy the lateritic hills of the
wandoo zone between Pingelly and Bavensthoi’pe, attaining their highest
development betw*een Pingelly and Wagin to the east and west of the Great
Southern Railway. Both species readily germinate after the country is
denuded by fire, forming dense thicket-hke areas over the lateritic hills, with
a shrubby undergrow'th in which Oxylobiurii parvifloruyn and Dryandra are
common, together with certain endemic species. To the south of Wagin, the
mallet areas occur on lower-lying ground, but not in depressions, wiiile the
two species extend in a mallee form almost to Esperance.

(d) FORMATIONS OF THE GRANITE ROCKS.

Throughout the interior, the basic granite is exposed in small rounded
hills or tors, often quite devoid of soil. These masses serve to collect amounts
of rain water w'hich runs off their sides, or collects in shallow pools on the
rock. The areas thus receive more soil moisture around their bases than the
surrounding country, and this is of importance to the vegetation. There
are a number of plants entirely confined to the granite rocks and their vicinity.
Amongst the trees, the most characteristic is perhaps Casuarina Huegeliana,
which attains a height of five or six metres, w'ith heavy pendulous branches.
Other characteristic species in the interior are Leptospermum ertibesce?is,
Kunzea sericea, a shrub with handsome scarlet flowers produced in great
abundance, and often growing out of crevices in the bare rook, and the two

The Vegetation of Western Australia.

xlvii.

ferns Pleurosorus rutifolius and Cheilanthes tenuifolia. Plagianthus Hehnsii
is perhaps more characteristic of the Eremaea, but characteristic of the granite
rock anywhere through the South-West Province and most of the southern
Eremea are the pin-grass {Borya nitida) and certain species of Drosera and
Orchidaceae. Certain south-western elements are, through the additional
moisture of the granite soils enabled to obtain a foothold far beyond their
usual climatic range, and thus we find Nuytsia floribunda’ growing as far inland
as Yorkrakine, while Hakea prostrata and Hypocalymma angustifolia may be
seen as far inland as Southern Cross. It is during the winter, however, when
the shallow moist soil underlaid by the impervious granite rock receives its
abundance of moisture that the granite rock flora is richest in species ; then
the ground is carpeted with a wealth of bulbous Liliaceae, Orchidaceae, Drosera^
Stylidium, and Polypompholyx^ together with certain ephemeral Compositao
and Centrolepidaceae. There are, in addition, a number of species quite
restricted to the granitic formations, such as species of Ricinocarpus, of Acacia
(e.g., A. restiaceay A. lasiocalyx)^ Eucalyptus, Melaleuca, and other Myrtaccae,
as well as species of Hydrocotyle, Anthocercis, Lobelia, and many others.

(e) SWAMP FORMATIONS.

Extensive alluvial formations occur at intervals on the coastal plain be-
tween the Hill River and Israelite Bay. In their most typical form they
occur on sandy country between the edge of the plateau and the coast, and
retain their surface moisture throughout the winter and frequently until the
end of the year or, on the other hand, they are perpetually moist, especially
in the southern areas of their distribution. While it is quite impossible here to
go into details, it may be said in general that these areas are rich in annual herbs
in contrast to the surrounding country, and that Myrtaceae and certain Pro-
teaceae make up most of the shrubby and arborescent species. The number
of endemics is not high when compared with other formations such as the heath.
Prominent amongst the trees are the species of Banksia, especially B. littoralis,
B. occidentalis, and B. sphaerocarpa. The Myrtaceous genera include several
species of Melaleuca, especially M. parviflora and M. raphiophylla, a number
of species of Agonis, one' of which (A, juniperina) attains to a height of 15
metres ; Leptospermum, Beaufortia, Astnrtea, Viminaria, Boronia, Dasypogon,
Baxteria, Cephalotus, Oxylobium, Conospermuni, and generally the greater
number of the Restionaceae and Cyperaceae. Tlie herbaceous plants com-
prise a number of annuals and bulbous or tuberous-rooted plants such as
Tribonanthes, certain Orchidaceae, Polypompholyx and Utricularia, Stylidium;
Levenhookia, Triglochin, Drosera, Villarsia, certain Compositae, especially
the Angianthinae, Epacridaceae, and Proteaceae (especially Franklandia,
Conospermuni, and Adenanthos), and certain Goodeniaceae, especially Dia-
spasis and Anthotium,

Mention was previously made to the distribution of the phyllodineous
Acacias, and the fact that two species inhabited tlie swampy areas. It is
also noteworthy that certain genera which are well tlevelojjed in the dry
areas, even in the far away Eremea, are represented in the alluvial and swamp
formations. I would specially mention Bossiaea, Lachnostachys, Anthotium,
Dampiera, Hakea, and Melaleuca. Bossiaea and Lachnostachys are as well
represented in the dry interior a.s in the swamps, or even more so, while Dam-
piera and Anthotium, both of which are typical of most swampy areas, are well
represented in the dry lateritic soils of the Eremea.

Charles Austix Gardner.

(f) HALOPHYTIC AND DUNE FORMATIONS.

The halophytic formations will be dealt with at greater length imder the
chapter on the Eremean Province, since these formations differ very slightly,
if at all, within Western Australia, and are more typical of inland localities.
The only distinctive lialophytic formations of the South-West Province are
the salt marshes which fringe the estuaries, and the littoral marshes in proximity
to the coast. These are fiats which are wet or inundated during the winter
months, but completely dry in summer, except under unusual conditions.
The most characteristic plants are those broadly classed as “ samphires,”
belong to the genera Arihrocnemum and Pachycornia, halophytic succulents
which also occur in the small saline depressions found constellated through
the country, and which predominate in such localities. Other Chenopodia-
ceae are Suaeda maritima, Samolus rej^tcns and S. Valerandi, Didymanthus
Roei and Atriplex hypoleucay and certain grasses such as Puccinellia and Pholi-
zirus.

The dune plants possess little that is distinctively Australian. It is on the
littoral dunes that we find so many palaeotropic groups and species which
belong more distinctively to the Eremea. Amongst the former are Festuca
and SpinifeXy CarpobrotuSy Atriplex cinerea, Pelargoniumy Nitraria Schoeberi,
Cakile maritima (which is surely a natural immigrant), Arctotis nivea, a more
recent immigrant, and certain Australian maritime dune species such as
Lepidospemm gladiatum, Scirpus yiodositSy Scaevola crassifoliay and Acacia
cylopis. It is interesting to find here, plants which more properly belong to the
Eremea, such as Callitris, Exocarpus sparteOy Eremophila glabray Alyxia buxi-
Jolia, and Santalum acuminatum. These are found in soils which are physically
or physiologically dry, and the relationship of the maritime dune vegetation
to the Eremea is most interesting, but on the other hand, especially under soil
and temperature conditions, the strand vegetation cannot be taken as typical
of the country w^hich it fringes. In the northern areas, that is, between the
Hill and Murchison Rivers, there is an extensive development of a littoral
formation of shrubs or small trees in which certain species of Acacia play an
important part. To the north the area is rich in lianes, and although the
shrubby plants are in close association, there is also a marked development
of herbaceous species. Here again is a marked admixture of plants which we
regard as being typically Eremean, e.g., GyrocarpuSy Zygophyllum, Jasminum,
Hibiscus, and Cienfugosia, or tropical (Aphanopetalum and Dioscorea).

(g) SAND HEATHS.

Wherever the soil is of a sandy nature, whether a loose detritus or the
more compacted yellow or red sand of the interior, lieath-like formations
predominate. These heaths vary from typical low heath in the humus soils
of the southern littoral, to the taller heaths of the sand of the savannah and
sclerojjhyllous woodland zones and the northern littoral. The shrubs usually
vwy from -5-] metre in height, and are often intermixed with small thickets
of the mallee type of Eucalyptus, Melaleuca, or several Proteaceae, but their
constitution varies considerably within the various climatic areas, those
of the wet districts often merging into swam}) formations, while those of
the interior and tlie north usually occupy only the high ground. The most
extensive areas of this formation occur between the Hill and Greenough
rivers, between the Hutt and Murchison rivers, between Tammin and Ben-
dering, and in the southern littoral between King George’s Sound and
Israelite Bay. In other j)laces the heath areas, intersected by thickets and
woodland, form a broad |)attern in the broad zone lying between the 500
and 175 mm. seasonal isohyets.

The Vegetation op Western Australia.

xlix.

The heath formation is the real home of tlie true autochthonous flora,
which is exhibited here in endless variety. The most representative families
are Epacridaceae, Haemodoraceae, Casuarinaceae, Proteaeeae, Myrtaceae,
Verbenaceae, Rhamnaceae, and Goodeniaceae. On its broad mosaic pattern
we are able to delineate with fair exactness the limits of the Epacridaceae,
and such genera as Anigozanthus, Conostijlis, Drosera, Isopogon, Petrophila,
Conospermum, Lamhertia, Banksia, the various genera of the Chamaelaucieae,
Kingia, Xanthorrhoea, CalectasiGy and a large number of plants which re-
present the autochthonous element.

Characteristic of tlie heath formations is the development of dew, even
in the summer. When the formations of tlie heavier soils are quite dry in
the early morning, a heavy moisture between the months of November and
March characterises the heaths, even to the limits of the South-West Pro-
vince. Another characteristic is the erect or erect-spreading ericoid type
of leaf, and this, together with the erect branching of the shrubs, causing
an open fabric of the constituents, results in a high insolation. Several plants
possess, as a protective measure, a strong development of corky tissue at
the base of the stem. This is particularly noticeable in several species of
Boronia and Eriostemon.

In contrast to the plants of the heavier soils, the heath shrubs possess
very deep rooting systems. It is not uncommon to find shrubs less than
15 cm. high with a root system extending to a depth of over 60 cm. Most
of the plants have a strong j)rjmary vertical with deep horizontal secondary
roots, the tap-root being as a rule remarkably strong, with a fibrous cortex.

While the contour of the heath vegetation remains more or less uniform,
a number of sj^ecies usually occur whicli rise much above the general level.
Several species of GrevUlea, Xylomelum angustijoliu7n, Haken, Banksia, and
Actinostrobus are familiar examples. They either occur in small groups,
or may be found in isolation, standing like sentinels on the heath profile.
Otherwise the heath is very uniform in tlie height of its component shrubs.
Some of the heatli species, especially the Rhamnaceae and the Epacridaceae
bloom before the advent of the winter rains, for example, I have seen Leu-
copogon and Cryptandra in full bloom at the end of April at the end of the
long dry summer. On the other hand, due perhaps to the beneficial influence
of the dews, certain heath species bloom in the height of summer. Verti-
cordia and Co7iosper7mtm are notable in this respect, and it should be re-
membered tliat the glorious Nuytsia which is so aptly named flo7'ihunda,
and which flings its golden wealth abroad in both coastal woodland and
sand-heath, is a true jisammophyte, and most abundant on the low open
heaths between Esperance and the Oldfield river.

Amongst the heath plants one observes as in no other formation except
the swamp-yate woodland, a remarkable development of plagiotropism.
The most notable examples are the extraordinary species of BaTiksia of the
series Prostratae which occur all along the southern heaths between King
George’s Sound and Cape Arid. Their subteri’anean stems radiate to a
diameter of over four metres, the isolated inflorescences and floral leaves
of some s]]ecies aj^pearing as isolated plants. Other notable examples are
the species of Eremophila, Lesche7}avltia, Astrolo7ua, Scaevola^ Hibbertia,
Acacia, Grevillea thrysoides and G. nudiflora, Dodonaea, and others, including
several species of Darwinia.

It is in the lieath formation that the species of the South-West Pro-
vince exhibit their greatest epharmonic convergence. Species of entirely
different families may a]q)ear so alike in form and structure as to be indis-

Charles Austin Oardn'er.

tiiiguisliable oxcojjt when in flower. The most iiotowortliy examples are
to })(' found in Hfbberlia microphylla and Leucopogon gibbofiits, between
Avavi(( and J/akea, or Acacia and Oreinllea.

Finally na^ntion sliould be made of the taller jdants whieh occur in
thicket formation between the sand-lieath and the sclerophyllous woodland.
As a nik? the two formations an' se[3ai‘ated by a transitional zone in which
tiu' me.lle(' Kncaljiptv.s takes an important part, or else there may be a close
stand of Melaleuca^ especially d/. ^mvinaUt. 'Phis latter species cannot be
said to 1)(‘ typical of any distinctive formation. It may occur fairly ex-
tensiv(‘Iy in tlu' granite soils, es[)eeially in areas when' the granite has been
kaolinized : it may sej;ara-te the heatli from the formations of [)sammophyti(‘;
halojiliytes, or it may stand as a buffer b('tween the woodland and the heath,
wlu'u its cliaracb'r clianges according to the nature of tlie soil. Typically
the iK'ath mergc's into a band of dwarf Melaleaca which receives here aiul
th(T(' a ft'w j)sammo|)hilotis siiecies of Fiicalyptus ; gradually the malices
become more prevalent, until finally from an association of mallees we pass
into the open woodland.

3. THE EREMEAN PROVINCE.

Th(' Krcmean Province is, floristically and ecologically, the most im-
poverished of the tliree provinces. Possessing no distinctive elements of
its own. its flora is deriv('d from elements of the neighbouring provinces,
chiefly from those groups sufficiently plastic to permit of epharinonic moulding.
It is thus a comj)aratively young flora in which the outstanding characteristics,
a marked evolutionary epharinonic convergence, has been made possible hy
the plastic nature of the indigenous species and their capacity to adapt them-
selves to a changcxl environrnc'nt. Its permanent elements are those sclero-
phyllous species which have to fight for existence against a hostile environ-
ment. wliile its herbaceous elements are either ephemeral, or markedly xero-
phytic in structure.

Jn its evolution the whole constitution of the Eremean flora is a picture
of acti\’e epharmosis, resulting in an area of remarkable uniformity of contour
and form which is in keeping with its climate. It extends in a practically
unchanged condition from the north-west coast to the Da.i'ling River, and
almost to the south coast, with but little change in its floristic composition,
and its formations are edaphic. rather than climatic, for climatically it is
only the increasing aridity exiR'riencetl as wo travel towards the arid lu'art
of tlu' continent that affects to any extent the density and stature of its com-
poiK'nt species.

An interc'sting feature of the Eremea is the part played by tlie psammo-
phytcs. Wherever the soil is sandy we find a return of the autochthonous
flora of the south-M’est, or at least its hardy psammophilous representatives.
This peculiai’ity (;an be studied in various localities. In the iH'd sand of the
Anketell district we find an “ island ” of south-w(>stcrn forms — -Labicheay
Thryptotncnc. Crypiandray Calythrix, OastTolohiumy Brachyscma. PityrodiUy
Loudonia, Monotaxis, Eriosiemon, and Xanthorrhoca, etc. The same thing
applic's at Comc>t Vale, where the Mulga country impinges on the heath ;
here are certain localised erukmiics, including ci'rtain Myrtaceous jilants of
the Tribe t^hamaelauciae, with Newcastlia and Lachnostachys. Even in the
north, between the Eortescue aiul Ashburton rivers wo find Verticordia grayidis,
Pileanihus, Pityrodia, Calothamnus, Comnicrsonia, Diplopeltis, Adriana, and
Calythrix, while Cyaiiostegia extends to Roebuck Ikiy. These isolated areas
of sand in v'hich the autochthonous flora is predominant, forming as it were
outposts of the south-western flora, suggest that it is in these place's tliat the

I

1

The Vegetation of Western Australia.

li.

old pan*Australian element is maintaining its last stand, or on the other hand,
and more probably, that it is in this environment that as an unmodified in-
vading element the autochthonous flora finds conditions suitable to its de-
velopment. I have previously pointed out that most of the plants of the
Autochthonous element are psammophytes.

On the other hand, the loamy and alluvial soils of the Eremea provide
the environment for the hardier species of the Palaeotropic Element. The
species of Sorghum, together with many other palaeotropic grasses, Acacia
Farnesiana, Clerodendron, the Cucurbitaceae, etc., have established them-
selves in regions far south of the Tropic of Capricorn. For example, Acacia
Farnesiana occurs as far south as the Murchison Kiver, and Ficus occurs as
a lithophyte as far south as Mount Margaret.

THE FORMATIONS.

(a) THE SCLEROPHYLLOUS WOODLAND.

The sclerophyllous woodland of the Eremea is the same in composition as
that of tlie marginal areas of the South-West Province, differing only in the
increased paucity of its undergrowth, and in the fioristic composition, as well
as in the importance of grass, which increases as warmer temperatures are ex-
perienced, while the increasing aridity results in greater numbers of Cheno-
podiaceae in the open spaces. A brief description of the formation has already
been given. It is sufficient to say here that the formation extends furthest
in saline and loamy soils, the most inland locality being found near the sliores
of Lake Darlot, where a Eucalyptus woodland of E. Duyidasi and E. Le-Soueffii
occurs in saline soil with a ground covering of Chenopodiaceae, Myoporaceae,
and Malvaceae, including the remarkable Plagianthus Helmsii.

(b) SALSOLACBOUS SHRUB STEPPE.

To the east of Israelite Bay and Balladonia, there is an extensive area
known as the Nullarbor Plain, the limestone soil of which supports a vegetation
averaging 50-100 cm. in height, consisting chiefly of Chenopodiaceae, Myopor-
um, Eremophila, and other low shrubs which are distinctly Eremean in cliarac-
ter. During wet periods a remarkable herbaceous growth cpiickly appears,
to die again when the soil moisture is exhausted. I have no first hand know-
ledge of the formation, but apparently it is developed extensively in South
Australia. Its ephemeral herbaceous species are mainly Compositae and a few
annual grasses.

(c) MULGA BUSH.

The mulga bush covers a greater part of Australia than any other single
formation. The conditions limiting its distribution to the south are only
imperfectly understood, but the line corresponds roughly to the Meyer Ratio
isopleth of 25 inches, and more approximately to the 80° F. mean isotherm
for January and February (the end of the dry season). There is also a certain
change in the soil which may be the result of climatic factors, and a certain
pluvio-thermal significance in this line, since to the south, with a slightly
increased rainfall, the wet season extends from March to August, while in the
mulga bush, the six consecutive wettest months are January to June. The
annual rainfall falls short of 950 points. Little more can be said of the climatic
control except that the mean temperature for the four wettest months is in
excess of 18° C., while for the sclerophylous woodland to the south it is less
than 18°. Unfortunately there are few meteorological stations within the
area, and only two near the boundary, and thus our knowledge concerning
the controlling climatic factors is very limited. The line of demarcation is,

11.

Charles Austin Gardner,

however, remarkably well defined. One crosses it at Goongarrie to the north
of Kalgoorlie ; it runs exactly past Mt. Singleton, very close to Mt. Jackson,
and it is intercepted again between Pindar and Wurarga, Exactly what
happens to the east of Goongarrie is not definitely known, but the southern
boundary reaches the coast at Shark Bay. Travelling towards the mulga
bush we find that the Eucalypt/us shrubs and trees become more and more
scarce, except in the sand, the species of Acacia become more prominent, and
finally we enter into a region where the genus becomes dominant, the species
being shrubs 3-7 metres high, with rigid glaucous phyllodes.

I have already mentioned the fact that the species of the Eremea ex-
hibit a marked epharmonic development. Acacia gives us a case in point.
The common jam tree of the south-western savannah woodland {Acacia
acuininata) has green phyllodes 7-25 cm. long and 4-7 mm. broad, in the
typical form ; in loamy soils it develops as a shrub with narrower phyllodes,
but still green and flat. As we a])]3roach the mulga country, this shrubby
form becomes more and more common in the loamy soils, its phyllodes
gradually becoming narrower with the increasing aridity, until finally, in
the mulga bush it merges into Acacia Burhittii, witli terete or only slightly
flattened phyllodes, and smaller flower spikes. This gradual transition
may be studied anywhere between Mullewa and Meekatharra ; at the former
station we have typical A. acuminata, and at Meekatharra in the loamy soil
of watercourses we have typical Acacia Burkittii. A similar graduation
may be seen between Eucalyptus p>yriformis and E. Kingsmillii. The same
holds true for a number of plants, and this is now'here more evident than in
the genus Eremophila, particularly between E. Gilesii and E. foliosissima ;
but numerous examples could be given, and the above should suffice to il-
lustrate the point. This epharmonic development is indeed carried out to
such an extent in the Mulga country that ultimately, amongst the woody
plants only two or three types of growth form and foliage remain. In general,
with but few exceptions, the broad-leaved plants are either covered with a
close indumentum of stellate or floccose hairs, or are heavily coated with
resin. The glabrous leaved shrubs have narrow erect or pendulous leaves
which all assume the same form, and the branching habit is much the same
throughout.

The Acacia species are of three types : Those with piimate leaves in-

habiting the watercourses ; those with green pungent })hyllodes of the
‘"Curara” type with a similar habitat ; those with erect or drooping rigid
glaucous phyllodes — ^the true “mulga” plants — and those with pendulous
flat green phyllodes, e.g., A. quadritnargvnea.

Next in importance to the species of Acacia are the species of Eremophila,
wliich number about one hundred. No other genus in the province exhibits
such diversity of form. The large handsome blossoms are short-lived, and
in many species the calyx becomes enlarged and coloured after anthesis.
While some conform in their growth form to the general plan, others are
widely branched with large resinous leaves. They remain of enormous
physiognomic importance within the formation, and occur mostly in the
sandy or stony soils, the genus being perhaps the most typical of all Eremean
genera.

The distribution of Eremophila is interesting. By far the greater number
of the species of this genus are found in M^estern Australia, and although
some are plants of the heavier soils, by far the greater number are psam-
mophytes or lithophilous. This together with the fact that the genus is
fairly well represented in the littoral areas of the South- A\'est, and also on
the south and western heaths, suggests that it has developed from this pro-

The Yegetatiox of Western Australia.

liii.

vince, or at any rate from a littoral source. Their development suggests
that as halophytes they have been e(juipped for the dry and saline tracts
of the Ercmea. Indeed, it may be said that they are principally developed
in the Eremea as psammophytes or halophytes.

Together with Eremophila, the genus Cassia assumes considerable physi-
ognomic importance in the Eremea, being developed most extensively in
the sandy areas. The development of an indumentum of felt has enabled
these plants to endure extreme aridity without any loss of leaf-surface, a
fact which is curious when it is considered that the only species of the genus
which has invaded the marginal areas of the South-\^’est Province, has its
leaves reduced to two or three terete segments. The desert species —
C. clesolata and C. SiMrtii — are examples of the plants which, endowed with
a thick felted indumentum, have, without any reduction in leaf-area been
not only able to exist here, but become increasingly important in the more
arid regions of the north, extending into the steppe formation.

Perhaps the most remarkable tree of the Mulga country, and of the
Eremea generally, is the Kurrajong {Brachychiton Gregorii). This tree, which
attains to a height of seven metres, has the spreading branches and dense
leafy crown of Platanus^ which it somewhat resembles ; its large bright green
broad leaves are partially or completely deciduous, and its trunk and roots
store water which is used in periods of excessive drought, at the time when
leaf-fall takes place. It is an alien aristocrat of the Eremea, cpiite unlike
any other growth form, and while of undoubted palaeotropic origin, has
extended as far south as temperate latitudes. It is still in evidence at Westonia,
Bullabulling, and at Lake Lefroy near Widgiemooltha.

The number of palaeotropic elements amongst the undergrowth of the
mulga bush is surprisingly large : examples are Breweria^ which extends

also into the northern parts of the South-West Province, Tecoma, Justicia,
Plectronia, Pomax, Hibiscus^ Gossypium, and amongst the lianes, Marsdenia,
Parana, and Pentatropis. They are found principally in the shade of the
larger species, often clustered round the stems (especially Ruellia), or in stony
soils.

Most, if not all of the woody elements of the mulga bush, possess seeds
remarkable for their longevity. The seeds of some species of Acacia, for ex-
ample, may lie in the soil in a viable condition for upwards of a century — a
factor of considerable importance when we consider the capricious nature
of the rainfall and the periodic desiccation by drought which characterises
the region. In addition, the species of Acacia, unlike those of more favoured
regions, produce their blossoms shortly after rain falls. Blossoming may occur
twice in the same year, or on the other hand, it may be withheld for a number
of years. There is thus no regular flowering season for the species of this
genus, while on the other hand, the species of Eremophila and Prostanthera,
Dodonaea, etc., appear to have prolonged regular seasons. In fact, the species
of Eremophila bear at least some flowers almost throughout the year.

The wealth of the ephemeral flora of the mulga country is one of the
most remarkable of its characteristics. The formation is typically one of
widely spaced shrubs with bare exposed soil, but after suitable winter pre-
cipitations the whole area is carpeted with an amazing wealth of annual herbs
which cover it with colour. These consist mainly of Compositae : wide

stretches of Helipterum splendidum and Cephalipterum Drummondii in the
alluvial soils appear from a distance like patches of freshly fallen snow ; the
sandy areas are carpeted with Cala?idrinia, Erodimi, the handsome rose-pink
Velleia rosea, purple Bellida, the gold of Helipterum Battii and H, Charsleyi,

liv.

Charles Austin Gardner.

and tlie violet and white of Calotis, Brachycome, and Erodiophyllum, while
the masses of the vetch-like Swainsonas intermingled with the whole provide
a herbaceous floral wealth unexcelled in any other formation. This brief
pageantry continues only for a few weeks ; two months later there remain only
the withered remains of stalks as evidence of this.

On the other hand, the summer rains call into being an almost equal rich-
ness of grasses and Trichinium, the former consisting mainly of short-lived
annuals such as Aristida and Chloris, Brachiaria, Setaria and a few others,
at the same time reviving the dormant stocks of the perennial species such as
Eragrostis, Neurachne, Triodia, Plectrachne, and Eriachne. Stipa, the prin-
cipal representative of the sclerophyllous woodland, is entirely absent, and
Amphipogon does not extend far into the formation from the south. The
mulga-bush formation exhibits this biannual succession to a degree not ob-
served in any other formation in Western Australia. In both cases it is
short-lived, without a sufficient rainfall it does not occur, and for years no
annual species may appear.

The true mulga (Acacia aneura) is the most common species in the mulga
bush, and its range extends throughout the formation,, or practically so.
It is most typically developed in those soils with a hard subsoil close to the
surface ; in the lighter deeper soils it becomes scarce or entirely disappears,
and its place is taken by Acacia linophylla in the area between the Minilya
and Murchison districts near the coast, while farther north, between the
Minilya and Fortescue Rivers, Acacia xiphophylla assumes a dominant role,
but even here, in the hard-pan soils, or in clay depressions, Acacia aneura
reappears, and once more becomes dominant to the east of Mount Bruce
in the upper Fortescue watershed.

Over very large areas of fiat or undulating country the Acacia species which
dominate the formation provide a uniformity of contour and physiognomy
to which the formation owes its name. The various species so closely resemble
each other even when in flower, that it is often impossible to determine ac-
curately the component species, unless pods and seeds are available. The
species are so plastic, that it is probable that ultimate research will disclose
fewer species than those already described, since the so-called species appear
to merge into each other. The landscape resembles a flat or rolling sea of a
dull grey-green, relieved only by the rocky eminences, and the scattered
salt-lakes which glisten in the sun.

The stony hills and ‘ ‘ breaka-ways ’ ’ provide some relief from this monotony,
for here we find a more varied flora. Here we find the bright green masses of
Dodonaea filifolia covered with crimson hop-like fruits, the green-leaved Plec-
tronia, Santalu7ni Hibiscus, Eriostemon^ Grevillea extorris, an occasional Melaleuca
or Tliryptomene, and the green-leaved Acacia quadrimarginea, mixed up with
brilliant masses of Eremophila, and the large-spiked Trichinimn rotundifolium,
•with several herbaceous species of the same genus, together with the dense
Cupressus-like masses of dull green Callitris glauca. The white corky barked
Trihulus platypterus, with its pale pinnate leaves and yellow star-like flowers
adds a bizarre note to this strange assembly in an environment so poor in forms.

(d) THE HALOPHYTIC FORMATIONS.

Throughout the Eremea, salt-pans, clay-pans, and the remains of former
ancient river systems are scattered. They form perhaps the most striking
physiographic feature of the landscape, appearing i ntensely bright because of
the reflected light of the salt crystals of their dry surfaces, because of the
remarkable mirage, and lastly because of the distinctive grey vegetation which

The Vegetation of Western Australia, i

Iv.

covers their broad margins. In this vegetation the Chenopodiaceae assume
the leading role. Species of Atriplex, Bassia, and Kochia cover the ground
with their lowly forms, relieved here and there by the more halophytic members
of other families, which form a striking contrast in colour — the green of Hakea
Preissii and Gasuarina lepidophloia, although the latter is green only by com-
parison, for its branchlets are more or less glaucous. The larger shrubs are
mainly species of Eremophila, such as E. pterocarpa and E. florihundai both
with dense rounded crowns of light grey foliage. Only where the soil has been
heaped into small rises do we find any relief from the drab colour scheme, and
here we find the purple-spiked Trichinium exaltatum, stretches of pink-flowered
PranJcenia, dense masses of halophytic Swainsona, with Senecio and Peplidiuniy
while in the south we occasionally find dense stretches of rose-flowered Carpo-
brotus. In the salt-pans of the mulga bush we find two halophytic Malvaceae —
Plagianthus microphyllous and the remarkable P. Helmsii, the latter with
a candelabra-like habit and densely packed clusters of leaves and flowers,
looking from the distance like a Euphorbia. The hardy halophytic grasses
are species of Eragrostis, especially E. falcata and E. lanipes, which intrude
sparingly into the area dominated by Bassia and Atriplex. The margin of
the surface salt, beyond which no plant life extends, is usually indicated by a
band of the green samphires which always remain as the last rampart of the
halophytic association.

(e) THE TRIODIA STEPPE.

Grass steppe occurs on the red sandy soils of the Eremean Province. It
is most typically developed in the areas which receive a summer rainfall,
but in a modified form it extends into the middle zone of the Eremea where the
highest precipitations occur between March and June. It is entirely absent
from the May-August rainfall zone except in a very impoverished condition
near Coolgardie, to the east of Mount Holland, and near the Fraser Range,
where the presence of Triodia tussocks in the sand-heath formation render it
peculiar. It is, however, exactly in these marginal sand-heaths that the
March rains are very important.

Speaking generally, the Triodia Steppe is a formation controlled by
summer precipitations and a sandy soil. It is the red sand that supplies
its peculiar edaphic requirement, except in the tropical north, where it occurs
in the stony soil of the Nullagine series, as for example the mountains and
escarpments of the Hamersley Ranges and the Hann Plateau in Kimberley.

The predominant genus is Triodia, comprising about a dozen species.
This grass forms dense tussock-like masses which usually average about a
metre in diameter, and may be cushion-like, dome-shaped, or conical. They
are very rarely contiguous, but are separated by narrow patches of bare sand,
and their growth continues peripherally, until in old plants the tussocks may
be as much as four metres in diameter, ultimately tending to become annular
as the centre dies out. Frequently, as a result of fire, these annular growths
are interrupted, and from small nuclei, fresh tussocks are again formed, which
in turn repeat the process. This is particularly true of Triodia Basedowii and
T. irritans in which, in the old plants, the progressive annular development
of the tussock results in some overlapping of the rings, causing a loose net-
work of living culms with dead centres, or the centres may be represented
by areas of bare red sand. The leaves of all species are pungent-pointed,
a characteristic that renders the plants forbidding.

The formation extends to the coast between Onslow and the Eighty-mile
Beach, but is otherwise absent from the littoral. From the north-west coast

VI.

Charles Austin Gardner.

it extends almost uninterruptedly through and around the Hamersley Range
and dominates the northern part of the Eremean Province between the Mulga
Bush formation and the savannah formations of the north. In some places
it intrudes into the Mulga Bush country w'here the red sand provides the
necessar^^ environment, but here it loses much of its character by reason of
the presence of the mulga trees and other shrubs. In its typical form, the
Triodia Steppe is either entirely devoid of trees and shrubs, or there may be
a fe-w isolated trees, such as Acacia pryifolia near the north-west coast, or
scattered Eucalyptus trees in the interior, or some admixture of Eremophila
and Cassia. There are large areas to the north-east of Wiluna, and east of
the Carnarvon Range, where, as far as the eye can see there is nothing but
Triodia^ unrelieved by any shrub or tree. It is only under the more favourable
conditions of higher rainfall towards the Northern Province, that trees become
relatively important, and the steppe formation merges into the savannah, and
other grasses compete with Triodia, e.g., Eulalia, Chrysopogon, and Eriachne.
Before reaching the Fitzroy River it merges into the Pindan country, in whicli
Acacia becomes predominant.

Just as to the north it takes on something of the characteristics of the
savannah and the Pindan, so to the south, in the Mulga formation, the in-
fluence of the winter rainfall causes some local alteration to the physiognomy
of the steppe by the addition of shrubs and trees, and the formation receives
to some degree the addition of the ephemeral winter flora wJiich characterises
the Mulga country. The steppe formation remains, however, a formation
restricted to the Nullagine rocks and the red sand, and nowliere is it found
away from this soil type, neither does it flourish in the absence of summer,
precipitations. Tlie drier the conditions, so much more does Triodia become
dominant, until it enters into the true desert.

(f) THE DESERT.

While, as its name would imply, the whole of the Eremea conforms with
the broader deflnition of what ecologists term desert, there is an area close
to the centre of Australia which complies with desert in its more limited
sense. The extreme aridity of the country, the absence of any permanent
surface water, the high annual mean temperature, and the extreme diurnal
range, as well as the paucity of the vegetation, are all conditions characteristic
of deserts in other parts of the wmrld. This tract, in Western Australia, lies
betv-een the latitudes of 20° and 25°, and extends eastwards of longitude 122^^
as far as longitude 132°.

For our accounts of this country we have to rely largely on the published
descriptions of the few intrepid explorers who have penetrated the region.
These accounts differ to some extent ; some state that there is no desert in
Australia, meaning that there are no large areas of bare sand ; others speak
of it in unmistakable terms. The fact that no botanical investigations ha^’e
been made, renders a true conception somewhat difficult.

In general, the region consists of large areas of sand ; sandstone cliffs
mark the escarpments of plateaux which are either sand or gibber-plains,
while there are depressions of sandy loamy soil. But it is the red sand that
prevails, either in the form of undulating country, or heaped into long ridges
or dunes which have in general a meridional trend, or bear in a N.N.M .-b.S.E.
direction. These dunes or ridges are either quite devoid of vegetation, or are
populated by sj arse Triodia and a few harsh xerophytic shrubs, while in the
trough-like depressions between tlie ridges there are sparse dwarfed trees and
shrubs ; on the other liand there are extensive undulating areas of bare wind-

The Vegetation of Western Australia.

Ivii.

eroded sand, and gibber-plains on wliich there is no vegetation of a permanent
kind. The only trees met with are in the dej^ressions or at the foot of the
escarpments, the most common being Hakea lorea, the desert shcoak {Casuar^
ina Decaisneana), Eucalyptus gamophylla, and E. setosa, the two last-named
rarely exceeding 10 or 12 feet in height. A few plants of mulga occur liere
and there in the depressions, e.g.. Acacia sibirica and A. Kempeana, while in
the heavier soils are a number of Chenopodiaceae, such as Kochia, Bassia^
and occasionally Atriplex. Following adequate precipitations, an ephemeral
flora is born, much resembling that of the savannah and Mulga-bush country,
but it disappears after completing its brief life cycle.

The sand ridges are usually scantily clothed with shrubs, amongst which
Crotalaria Cunninghamiiy NeiOcastlia, and Duhoisia Hopwoodii are the most
common, while on the flat sandy intervening country are occasional shrubs
of Hakea rhomhalis, Orevillea eriostachya and G. juncifolia, Calothamnus, Dicra-
styles, Thryptomene Maisonneuvii, Dampiera, Petalostyles, Hibiscus pinonianvs,
Brachysema Chamhersii, Sida and Acacia, Cassia desolata, and Micromyrtus.

Tt is interesting to find here such plants as Dampiera, Brachysema, Colo-
thamnus, Thryptomene, and Micro^nyrtus. They are all local endemics derived
from a south-western stock, and are psammophytes equipped for a hostile
environment. Crotalaria, Hibiscus, and Petolostylis are of northern origin,
and their appearance may seem strange, were it not for the fact that all possess
an indumentum which enables them to thrive far removed from their main
centres of distribution. One should perhaps in this connection mention
Gossypium Sturtii, although we do not know whether this species is a psammo-
})hyte, or confined to rocky gullies.

The only endemic genus of the true desert is Newcastlia, which through
Hemiphora, Lachnostachys, and Dicrastyles exhibits a certain aftinity with
the South-West Province, and once again illustrates the fact that it is the
psammophilous elements of the south-west which have not only exceeded the
boundaries of the South-West Province, but have been able to create new
forms in an arid environment.

As the gradual process of continental desiccation progresses, so also is
the desert extending. Under the influence of the prevailing winds it is grad-
ually encroaching upon its neighbouring formations, thrusting here and there
into the Triodia Steppe, and while it is extending to the coast in the vicinity
of the Eighty-mile Beach, another arm of the steppe is extending along the
Tropic of Capricorn to the coast, in the vicinity of the Ashburton Kiver.
There can be little doubt that the Hamersley district is threatened with an
Eromean invasion which will, sooner or later, completely overrun it.

Vlll.— EPHARMOSIS AND GROWTH FORMS.

Just as the species is the unit of the systematic botanist, so is the vege-
tation form” the unit of the ecologist. To quote Warming : “ Every species
must be in harmony, as regards both its external and internal construction,
with the natural conditions under which it lives ; and when these undergo
a cliange to which it cannot adapt itself, it will be expelled by other species
or exterminated. Consec^uently, one of the most weighty matters of
ecological plant-geography is to gain an understanding of tlie e])harmony
of sj)ecies. This may be termed the plant’s growth form in contradistinction
to its systematic form.” It reveals itself specially in the habit and in the
form and duration of the nutritive organs, but shows to a less extent in the
reproductive organs.”

Iviii.

Charles Austin Gardner.

A well-known result of epharmosis is the convergence of systematically
removed types. Exam]des are to be found in certain plants, such as be-
tween Ceriops and j^giceros, Accria ard Hclea, Acacia and Lavie^ia,
Darwinia and Pintelea, Siegfriedia and Durwinia, Hihbertia and J^eiicopoyon,
Hihhei'tia and Piittevaea, of wliich some species of one genus so closely resemble
species of the other genus that, growing together they can only be se|:arated
when in llower. The careful climatic grading of the Western Australian
flora has resulted in numerous epharmonically moulded species and forms
which attain to their highest de\elopment in tlie Eiemea where the ephar-
monic convergence reaches a climax in which the species conforni to few
general growth-form types.

GROWTH FORMS.

1. TREES AND SHRUBS.

Trees and shrubs make U]) the greater part of the vegetation of WVstern
Australia. Their importance is, however, greater in those areas dominated
by the winter rainfall- — summer drought climate than in those areas in which
summer rainfall prevails. It is especially true of the Soiith-West Province
where the ])re]3onderance of shrubs is overwlielmingly great when comj'ared
even to trees.

One of the outstanding characteristics of the forest and woodland
formations of South-AVestern Australia is the paucity of the tree species
growing in intimate association, with one of them assuming tlie dominant
role. This is particularly true of the karri, jarrah, and tuart forests where,
subject to edaphic changes, the dominant tree suffers few or no competitors.
For example, the tuart forest remains essentially jDure ; the jarrah tree
almost so, except where the lateritic soil changes to sand, marri (EucaJyj)tus
calophylla) assiimes prominence, while clay soils give rise to trees of wandoo.
Tliere is, perliaps, nothing comparable to this uniformity of the s]X‘(-ies of
the Eucalyptus forests and w'oodlands if we except the coniferous forests
of tlie northern liemisidiere. The savannah (temperate) M^oodlantl form-
ation is more mixed in its arborescent species, and Acacia acuminata, Eu-
calyptus loxophleba and E. redunca var. elota become social species. It is
only in the eastern sclerophyllous woodland areas that we find any real
mixture of the tree species, and although all are species of Eucalyptus, we
find a considerable mixture of species. In the Eastern goldfields areas we
find many such examples : the trees may very closely resemble eacli otlier,

such as E. Flocktoniae, E. BroeJewayi, E. transcontinentalis, E. salmonophloia,
which are very much alike both in habit, bark, and foliage, or dissimilar
species may grow' in intimate association. Between AVhdgiemooltha and
Norseman, for examj)le, or between Mount Holland and the Bremer Range,
one frecjuently finds as many as fifteen different species growing in asso-
ciation under similar edaphic conditions.

In the northern Province it is fjuite otherwise, for in the monsoon wood-
land we find trees of different families, of very diversified growth-forms,
even deciduous and evergreen, growing together. It is jjossible to find
Callitris, Eucalyptus, ami Livistona assuming ecpial importance and growing
together in a woodland association.

While Eucalyptus, remains dominant in the South-AA^est, Acacia assumes
the dominant role over the broad area of the Eremea, and in the Northern
Province, especialh' in the Kimberley district, there is no dominant genus
amongst the trees if we except the basaltic savannah, w'liere Eucalyptus
becomes again dominant. The Eucalpyius trees of the north how'ever, remain
highly diversified in both growth form and foliage. The dense spreading

Thk Vegetatiox of Westerx Alt.stkalia.

lix.

crown and large horizontal leaves of Eucalyptus clavigera and E. setosa^ for
example, are quite unlike those of any Eucalyptus of temperate latitudes, and
closely resemble Afigophora, The same may be said of the curious EucalyjHus
brarhyandra with its widely spreading branches and deciduous foliage.

Quite otherwise is the state of things in the South-West Province and
the Eremea. Here we find amongst the species of Eucalyptus much the
same leaf form, and, with the exception of Eucalyptus calopdrylla, E. ficifolia,
and E. haeniaioxylon, trees with the same shape of leaves, and a light crown.
In the C'omparatively wet forest areas there is development of spreading
branches, but when v'e travel eastwards to the woodland formations we
find, first typically exhibited by the swamp yate and the York gum, a growth
form whicJi becomes the dominant form throughout the savannah wood-
land, the sclerophyllous woodland, and the mulga bush. Not only is this
form common to the trees, but it becomes important in the shrubs also,
it is ]:erhaps most typically represented by the jam tree {Acacia acuminata),
but is so consistent both in Acacm and Eucalyptus throughout the drier areas,
that a description of Acacia acuminata will suffice. In this growth form,
which 1 shall lefer to as the Infundibular growth form, the stem, by a system
of repeated upward branching, produces a multiplicity of branches and
twigs, all attaining more or less the same height, and resulting in a fiat or
convex dense crown with a light covering of leaves. Seen in lateral profile
it resembles a funnel with a convex apex, the branches and branchlets leaf-
less except at the summit. Sometimes the crown is proportionately small
compared with the height of the tree, especially in large morrel trees (A\
longicornis) in the eastern districts. Whereas in Eucalyptus tlie foliage is
pendulous, in Acacia it is usually stiff and erect.

Amongst the shrubs we find the same infundibular growth form in many
plants, notably in Myrtaceae and Myoporaceae, e.g., Melaleuca uncinata, Ere-
mophila interstans and its allies, and in Verticordia. Verticordia Brownii offers
an excellent example. This plant, knovm as the “ cauliflower bush,” has
its dense convex surface so closely packed with white blossoms that no inter-
stices are formed, and no leaves are visible. The axillary opposite branchlets
are richly developed, resulting in annually produced dense strata of leafy
shoots which, viewed laterally, appear as a series of superimposed dense
umbrella-like corymbs separated by longer intemodes. The flowers are
densely congested in the uppermost axils, forming a compact floral mass ;
the leaves subtend the pedicels, and usually persist for a second or third vear,
thus providing a leafy substratum. The richness of the branching is remark-
able, and the lower strata consist of the dead remains of branches, the further
growth of which has been prevented by the overshadowing of the terminal
canopy. The whole resembles an umbrella in which the surface is annually
renewed, and each year's successive vegetative growth is plainly discernible.

As an adaptation to withstand the desiccating action of dry winds, this
growth form is obviously most effective, since only a minimum of the leaf^' area
is subjected to the action of such winds. It is associated with an erect, or
pendulous foliage, so that a great number of leaves can be carried without over-
lapping. It is usually the uppermost stratum of the crown which carries
leaves, and a superimposition of the leafy crown is very rare. That this
habit is not due to photophily, as has been claimed, is suggested from the
fact that it is most highly developed in the Eremea on the loamy soils where
the j^iants are widely spaced. This growth form is 2 )rcdominant in the Eremea,
where sometimes the epharmonic convergence is carried out to the extent that
it is often impossible to distinguish jjlants of different families when not in
blossom ; it is not developed in those j)lants wiiich possess an indumentum of

lx.

Charles Austin- Gardner.

felt or hair, or only to a very slight extent. It reaches its epharmonic climax
when complete aphylly is secured, as in Casuarina and Acacia, and the genista-
like forms of Eremophila scoparia and Calycopeplus Helmsii.

Occasionally in the mulga-bush we find a different growth-form adapted to
withstand the action of dry winds. This is the ereot -stemmed, horizontally
branched form which is typical of Acacias of the A, Grashyi and A. sihirica
type. The simple stem is widely branched with horizontal branches which
are again repeatedly horizontally branched to form fiat superimposed layers.
The lateral outline is thus a series of horizontal planes of leafy twigs bearing
short erect pine-like foliage. It employs less w’ood than the infundibular type
because of the superimposition of permanent leafy strata, and it is curious
that it should be so rare in trees and shrubs of the arid areas. In both this
and the infundibular form, the branchlets are abbreviated, a character common
to all the w^oody plants of the dry areas.

Wliile the infundibular grow’th form leads ultimately to leaf-reduction
or even complete aphylly in its epharmonic development, the widely branched
plants of the Eremea retain their leaves in an unmodified condition. The
I^inoid leaves usually arise from abbreviated axes and are thus fasciculated,
while the broad-leaved plants have relatively long internodes. Such plants
retain their comparatively large leaf area by the production of a protective
covering of felt-like hairs, densely interlocked stellate hairs, or the provision
of a viscid superficial layer of resinous material. No genus so typically affords
examples of all types as does the genus Eremophila. In the section Pholidia
w’e have ericoid leaves ; in Eremocosmos there is a tendency for the branchlets
to assume the function of leaves, resulting almost in complete aphylly in
E. Dempsteri ; in the -Sect. Eriocalyx we have flat leaves covered with felted
hairs, e.g., E. leucophylla, while in Platychilus we have large leaves covered with
a lacquer of resinous material, as in E. viscida and E. Fraseri, while in the
Section Stenochilus all types are represented. A similar condition may be
seen also in Cassia, C. Chatelainiana, C. phyllodinia and C. eremophila, C.
desolata, and C. glutmosa respectively exhibiting the same forms.

There is one outstanding example in the Eremea of a tree that does not
conform to the general epharmonic pattern. This is the Kurrajong (Brachy-
chiton Gregorii), wdiich maintains a dense bushy crown of bright green soft
leaves which appear to be unaffected by climatic extremes. In this respect
it stands in the Eremea as an alien whose relatives populate the monsoon
wnodland, and its appearance here is certainly strange. It owes its capacity
to withstand the aridity because of its water-storing trunk, and its ability
to become deciduous in adverse seasons, being the only tree of its kind within
tlie Eremean Province.

The Proteaceae stand in striking contrast to the Myrtaceae and Mimosa-
ceae in that their growdh form is never infundibular, and rarely even corymbose.
Isopogon and Petrophila, for example, produce their new shoots close under the
terminal inflorescence, resulting in a di- or trichotomous system, the old
terminal infructescences becoming lateral, or remaining in the forks of the
branches. Occasionally the growdh is sympodially developed. Banksia and
■ Dryandra also illustrate this form.

In distinction to this is the remarkable further development of the stem
or branch found m certain iVIyrtaceae. In Callisteyyioyi, Beaufortia, Calo-
thamnus, and Regelia, for example, the inflorescence is apparently terminal,
but the rhachis of the spike continues the vegetative growth w'hich is often
already w^ell developed before flowering is completed, and branching usually
occurs imyyiediately above the inflorescence, and is usually verticillate ; the

The Vegetation of Western Australia.

\xi.

successive annual shoots can be ascertained by the number of the successive
infructescences .

The production of short lateral leafy shoots at the nodes is a characteristic
growth form of several Myrtaceae, Dilleniaceae, and Proteaceae, especially
in Hibbertia and Dryandra. This leads to the formation of slender elongated
branches or stems densely crowded with abbreviated leafy shoots often bearing
terminal flowers. It is seen to advantage in Hakea ruscifolia, in Hibbertia
teretifolia^ and in some species of Dryandra.

Finally, there are the chamaephytes whicli develop a marked plagio-
tropism in their growth. These are mainly the plants W'hich rise but little
above the soil, or are so closely pressed against it that they form dense mat-
like or cushion-like growths. They are typifled by Astroloma, Dodonaea Acacia
pulvhiiformis, Leschenaidtia Jormosa and L. Hibiflora, Scaevola. pulvinaris,
and species of Dampiera in the wet clay or sandy-clay soils of the southern wood-
lands in which Eucalyptus occidentalis is the dominant tree. On the sand-
heaths of the southern littoral occur Leschenaultia, Scaevola, Eremophila and
Banksia. In the clay soils of the areas between Norseman and Grasspatch
many examples occur in Acacia and Eremophila, while to the north the chamae-
phytes are perhaps best developed in the sandy or lateritic areas, Leschenaultia
macraniha and species of Darwinia, together with Actinostrobus acuminatus
being typical examples. In the case of Banksia repens, B. prostrata, and B.
Goodii, the stems are usually covered by the sand, and the isolated flower-
clusters in B. Goodii appear as isolated plants scattered over a diameter of
three or four metres which represents the extent of the branching system.
The inclusion of E^'emophila amongst the chamaephytes once more illustrates
the remarkable diversity exhibited by the plants of this genus both epharmoni-
cally and floristically. No other genus in \Ahstern Australia is more worthy
of study from almost every point of view within the study both of ecology
and systematic botany.

2. LIANES.

Using the term in its widest sense, the lianes occur in almost all forma-
tions in Western Australia, but their number is not large. As one would expect,
they are most common in the forest and littoral thicket areas, where Clematis,
Keyinedya, and Hardenbergia offer the best examples, and almost complete
the total list. On the western plain we encounter in the richer soils three
jjlants which are remarkable because of their restricted occurrence : Aphano-
petalum, from the Chapman River, Clematocissus which extends from the
Murchison River to the Hill River, and becomes leafless during the dry season,
and finally Dioscorea. Their occurrence here suggests that they are migrants
from the north, but their absence to the lun'th of the Murchison River is re-
markable. In the Northern Province the lianes are mainly recruited from the
Asclepiadaceae, Apocyriaceae , Papilionaceae, and Convolvulaceae. Marsdenia
australis extends as far south as the southern confines of the mulga-bush
formation, while Glycine extends much further south, associated with the
granite rocks. The other important lianes are to be found amongst the
Pittosporaceae, Billardiera, Marianihus, and Bollya, having an extensive
distribution through the South-West Province ; Marianthus erubescens and
Billardiera alone extending to the boundaries of the Eremea. Coinesperma,
with two twining species, almost completes the list of southern forms which
extend into the interior, although Kennedya jyrorepens is the solo Eremean
form which extends as a psammophyte into the desert. Cassytha has a general
distribution throughout Western Australia, but is more abundant in coastal
than in inland situations. It is almost absent from tlie far interior, but
common in the littoral thickets and the swamps.

Xll.

Charles Austin- Gardner.

3. HERBS.

The perennial herbs are not richly developed in south-western Australia,
except amongst the Liliaceae, Haemodoraceae, Droseraceae, Stylidiaceae,
Goodeniaceae, Cyperaceae, Restionaceae, and Gramineae. They are mainly
characterised by a caespitose or cushion-like habit of growth, or are provided
with strong rhizomes, tubers, or bulbs, and are mainly inhabitants of the
swamp formations, the granite rocks, or the savaimah woodland. The Orchid-
aceae in general also conform to this general requirement.

An exception to this general rule is to be found amongst the Stylidiaceae,
Goodeniaceae, and more particularly amongst the autochthonous Haemodor-
aceae, which latter especially are mainly psammophytes. Examples are
Anigozanthos, Blancoa, Macropidia, Conostylis, and Phlebocarya. All of
tliese possess strong rhizomes and have either narrow persistent sub-distichous
leaves, or their leafy parts wither during the dry summer. A further character-
istic is the hairy or woolly indument of their inflorescences, and sometimes of
their leaves also. The bizarre colouring of Anigozanthos and Macropidia has
made them famous.

The Liliaceae and Droseraceae on the other hand liave hypogeal resting
organs, usually corms or bulbs. While their development is fairly high in the
south-west, it is relatively low when contrasted with the floras of other lands
with a similar climate. It is indeed far short of what one would expect when
the almost similar conditions which are experienced in the Cape Province of
South Africa are considered. The most diversified habit is exhibited by
Drosera, which has erect branching forms in D. gigantea, rosette-like forms
in LamrolepiSy and several scandent species in Ergalenmi. The Liliaceae are
both bulbous and fibrous-rooted, or some, like Stypayidra and Arnocrinum
have strong rhizomes.

The Cyperaceae are mostly swamp plants, and attain their greatest de-
velopment in the lower south-west. A few venture into the sand heaths as
hard tussocky species, but they are entirely absent from the Eremea, except
as inhabitants of river banks or marshes. They are particularly well de-
veloped in the swamp lands of Kimberley.

The Restionaceae, unlike the Cyperaceae, are mainly psammophilous,
and are equally at home in the periodically swampy areas, or on the dry sand,
where they sometimes assume a position of importance. Most of them are
strictly endemic in the Soiith-West Province, but a few, such as Ecdeiocolea,
Lepidoholus, and Chaetanthus are more typical of the heath formations ex-
tending into the Eremea. The fact that the dry sand of the interior has
produced forms such as this, together with the locally endemic Hopkinsia
and Harperia, illustrates once again the remarkable psammophilous preference
of the autochthonous elements.

In the Eremean Province the Compositae become increasingly important,
and have been discussed under the cliapter dealing with the Mulga Bush.
Cala7idrinia, the “ parakeelya ” of the interior, is another example of
a tuberous-rooted perennial which thrives under dry conditions. The species
of Calayidrinia are of wide occurrence in the Eremea, and in addition to having
tubers, the leaves are succulent. Some flow^er when in leaf ; other, like C.
scldstorrhiza, commence to bloom after the leaves have withered, their delicate
l^lossoins defying the dry heat of early summer.

The grasses are of most importance in the areas which receive summer
rain. Grass predominates in most of the Kimberley and northern formations,
but is also important through a wide area of the Eremea extending south

The Vegetation of Western Australia.

Ixiii.

to Meekatharra and Sandstone, in the area which receives as a rule some
rain from the storms which visit the north-west coast. They are mostly
of the harsh tussocky type in the dry regions, but range from the soft-textured
species of the black soil plains to the uninviting Triodia and Plectrachne.

The perennial grasses are dominant in the savamiah, tlie savaimah wood-
land and the steppe. The commonest genera of the tropical savannahs are
Themeda. Gymhopogon, Iseilema, Astrebla, several genera of the Andro-
pogoneae, including Eulalia, Chrysopogon, Bothriochloa, and Dicanthium,
w'hich flourish in the red loamy soils, to the many species of the Paniceae and
the soft-leaved Triodias of the lighter soils. Sorghum stipoideum, the tallest
of the grasses is important in Kimberley, in the monsoon woodland, and the
sandstone savannah, extending also to the Pintlan near Broome. It attains
a height of four metres, and frequently possesses prop-like roots from its
lower nodes.

An examination of the tropical genera and species discloses the fact
that a number of these are native both to equatorial Africa and southern
Asia, e,g., Sehima nervosum, and a nulnber of others. The number of endemic
genera is very small.

The grasslands reach their maximum development on the loamy soils of
tlie northern rivers — tlie “ black-soil plains ” — alhivial areas in whicli the
soil is rich in humus. Such areas are to be found on the Lemnard, Fitzroy,
King Edward, and Drysdale rivers, as well as the Ord River, while there is
an extensive area between the Prince Regent and Glenelg rivers. Further
south, in the Fortescue-De Grey district, Eragrostis, Eriachne, Enneapogon,
and Aristida become more important amongst the perennial species, forming
in places large areas of open grassland on the alluvial soils. These grasses
either occur as scattered dwarf caespitose clumps in the dry season, where,
possessed of strong rhizomes and a dense protection of crowded outer dead
leaves they are able to survive sustained periods of drought, or the small
stumpy base of the plant is the only evidence of life. With suitable summer
rains, liowever, they develop into a dense ground covering intermixed with
shorter-lived grasses and other herbs. I have not seen this country under
these favourable conditions, but from the impoverished residuum seen in dry
years, and the development in the depressions locally known as “ crab-holes,”
there can be little doubt that under conditions of adequate seasonal rains,
these areas present a picture comparable with the South African veldt.

The hardiest of all the grasses is Triodia, the “ spinifex ” of the interior,
which is the most important element of the steppe, and which is also at home
in the desert. Triodia must be regarded essentially as a psammophyte, for it
is never seen in loamy soils, where its place is taken by Eragrostis, Eriachne,
or Enneapogon, Not only has it a general distribution in sand throughout
the north, but it has to a great extent overcome climatic barriers, and has
extended as far south as Coolgardie, Mount Holland, and the Fraser Range.
It is, however, only in the stony or sandy soils of the Nullagine series tliat
it is n'ally at home. Few other grasses are similarly constructed to with-
stand the climatic extremes it has to endure, and it thrives under conditions too
dry for most other plants. The commonest speeaes of the dry steppe, it is
also an impoi’tant constituent of the mulga bush to the e^ast and north, anel
its prevalence or absence to a great extern! eletermines the limits of the steppe
formation. It always occurs in large scattered tussocks, and by a peripheral
system of stolons or rhizomes incrc'ases its girth until it ultimately extends

!xiv.

Charles Austin Gardner.

over a diameter of over tive metres. Fire destroys the aerial parts, and
from the periphery a ring-like growtli ensues, or this may be broken up into
a number of independent plants.

The distribution of the grasses in Western Australia is interesting, sineo
it shows that they fall mainly into tw'o thermal groups — a megathermic group
of (>3 genera, representing mainly the Paniceae, Andropogoneae, and I'lra-
grosteae, with a number of endemic genera, and a mesothermic group of 17
genera, represented chiefly by the Festueeae, Aveneae, and Agrosteac, with
but one endemic genus, Diplo 2 :>ogon. There are few genera which are common
to both areas^ — Neurach^^e extends to the north, but is more proptu'ly to be
considered of southern derivation ; Triodia, Eriachne, Eragrostis, and Sporo-
bolus extend far to the south, but must be considered a northern element >
The Sections Arundineae, Chlorideae, Zoiseae, and Arundinelleae are ox-
clusively groups requiring a warm summer rainfall, and the Andropogoneae
are characteristically of this element also. In the Stipeae, Aristida is mega-
thermic, while Stipa, its southern counterpart, is mesothermic. Stipa and
Aristida are common only to the eastern districts, the former being mainly
south-western, the latter northern.

It is imj)ortant to consider also the recpiirements and the development
of the grass flora in south-western Australia. The grasses here are sometimes
of a very restricted habitat. For example, Tetrarrhenn and Microlaena are
confined to the forests and woodlands of the highei* winter rainfall areas ;
iSporobolus, togetlier with Puccinellia, Ayuphibro^nus, Deyeuxia, Poa, Era-
grostis, Lep>turus, PholiurtiSy and Dichelachne show a j^reference for loamy or
swampy soils, wet at least in winter ; while Dantkonia, Neurachne, A)nj)hi-
pogon, Diplopogon, and Stipa favour the sandy soils, and attain their
greatest development in the southern regions whei’e the summer rainfall
is higher than elsewhere. For example, Danthonia is not common on the
west coast further north than Cingin, while it is fairly common along the
southern littoral. Festuca finds a suitable environment both on the littoral
dunes and the hard loam of the interior.

Apart from the shade-loving or paludal species, the grasst's of the south-
west appear to fa\'our the granitic soils, and prt'dominate in the savannah
woodland which separates the sclerophyllous woodland from the forest. The\*
are most abundant where the summer rainfall is higher than elsewhere (ex-
cluding the forest area), and under its influence we occasionally find an inter-
mixing of both northern and southern types, as at Jarramongup and C.'ool-
gardie.

In conclusion I must express my thanks to the Divisional Meteorologist,
Mr, A. G. Akeroyd, for permission to publish rainfall and other meteorological
statistics, also for kindly reviewing the chapter dealing with the climate ; to
Professor E. de C. Clarke for assistance in the chapter on geology and soils ;
and to Mr. F, M. Bennett for compiling some of the tables which ha\-e app(‘ared
in the appendix. Finally, I am indebted to Dr. Gentilli of the University
of Western Australia, for permission to use his statistics dealing with annual
rainfall variability.

The Vegetation of Western Australia.

Ixv.

LITERATURE.

Bentham, G. : “ Flora Aiistraliensis ” (1863-1878).

Bentham and Hooker : “ Genera plantarum ” (1872-1883).

Diels, L. : “ Die Pflanzenwelt von West-Australien ” (1906).

Gardner, C. A. : “ The Forest Formations of Western Australia.” Australian Forestry

Journal, v. vi, vii, viii, and x.

Gardner, C. A. : Botanical Notes, Kimberley Division. Bull. Forests Dept. (1923).
Hooker, J. D. : “ Introduction to the Flora of Tasmania ” (1859).

Jutson, J. T. : “ The Physiography of Western Australia.” Bull. 95, Geological Survey of
Western Australia (1934).

Warming, E. : “ (Ecology of Plants ” (English translation) (1909).

Ixvi.

Charles ArsTiN Gardner.

CLIMATIC TABLES.

)

TABLE I.— MEAN MONTHLY KAINFALL,

(In Points.)

The figures are complete to the end of 1941.

The figures in brackets represent the number of years during which records have been taken.
Stations in italics are those recording temperature.

The figures in heavy type represent the six consecutive wettest months.

l._XORTHERX PROVINCE.

Station.

Years.

Jan.

Feb.

Mar.

Apl.

May.

June.

July

Aug.

Sept.

Oct.

Nov.

Dec.

Total.'

Port George IV.

(27)

1,512

1,113

1,001

214

96

75

30

4

6

39

184

765

5,939'

Mount Barnett

857

701

421

103

21

11

19

9

18

96

419

782

3.457 i

Kimberley Downs

763

573

416

140

35

25

14

3

2

21

96

50

2.194

Beagle Bay

646

658

460

233

51

60

41

19

9

14

108

479

2,778

Turkey Creek ....

(44)

664

672

411

94

23

30

25

9

17

75

199

495

2,714

W'yndha.m

(55)

739

606

467

82

20

16

16

3

8

44

192

417

2,61(^

Derbu ....

C>6)

735

556

443

142

73

52

24

9

1

6

83

406

2,530

Broom p ....

(52)

648

568

393

115

61

96

20

11

5

3

56

325

2.301

Fitzroy Crossing

(48)

641

512

333

65

29

37

32

5

5

20

90

341

2.110

Hall’s Crepk

(51)

543

435

291

70

36

26

25

9

15

53

136

322

1,061

La Grange

(51)

435

436

413

95

82

92

32

9

3

3

37

233

] ,870

-\nna Plains ....

(34)

342

252

317

117

88

70

29

10

2

4

31

185

1,447

Wollal *

(32)

328

265

239

92

56

97

47

c8

4

3

11

146

1.206 ■.

Marble Bar

(47)

283

272

221

96

69

104

50

20

3

21

34

142

1.315'

Nnllaainp

(44)

307

219

218

83

66

SO

40

27

5

18

60

165

1 .291*

Hamersley f •

(14)

334

250

318

63

57

74

26

21

14

32

67

81

1.337;

Condon +

(37)

219

249

303

108

73

106

37

18

3

4

7

65

1,191 1

Port HpfUan'i ....

(44)

177

221

324

105

113

126

41

39

5

7

2

33

1.193

Itoebourne

(55)

202

196

293

140

100

102

57

25

7

3

7

36

1,168

Tambrey

255

357

346

113

97

129

64

35

15

8

31

112

1 .562

Millstream

270

316

288

127

92

136

69

27

12

4

31

105

1.477

Whim Creek ....

(43)

195

277

430

191

98

111

58

26

10

11

19

41

1.467

Red Hill§

(27)

262

309

228

110

83

147

70

29

10

7

19

73

1 .356

Bonnv Downs

(35)

237

211

189

68

76

76

32

18

6

22

61

136

1.132

Vlaming Head

123

261

208

160

108

247

49

49

5

1

4

0

1,215

Bamboo Creek

292

226

206

128

78

137

62

28

3

17

47

127

1,441

• Wollal recon

for 32 years ending 1918.

t Hamer.slej

records for

14 years

ending 1925

t Condon fl

reconls for 37 year

ending 1927.

§ Red- Hill reeords for 27 years endinc 1925.

ir.—EREMEAN PROVINCE.

(a) Sortherti Zone (Summer rainfall, Dec.-Ai)ril).

Station.

Years.

Jan.

Feb.

Mar.

Apl.

May.

June.

July.

Aug.

Sept.

Oct.

Nov.

Dec.

TutaL

Miindiu'indi

.... (26)

142

157

204

81

66

75

25

31

16

50

45

129

1.021

Ethel (’reek

.... (31)

170

185

193

87

66

78

27

11

15

20

44

96

1,001

Roy Hill

.... (42)

162

204

191

90

63

63

36

25

7

17

33

110

1,001

Three Rivers ....

.... (27)

125

88

178

75

114

83

36

17

8

20

37

91

872

Mardie . ..

.... (54)

120

147

171

67

120

115

52

38

7

4

3

26

8<0

Winning Pool ....

.... (44)

135

166

151

60

129

158

83

48

8

7

7

34

086

Wandagee

103

166

96

71

106

158

104

36

14

7

6

34

901

Peak Hill

(41)

124

112

170

92

97

116

55

40

17

14

35

68

940

Wiluna

.... (43)

132

105

142

135

98

84

47

37

17

26

31

84

938

Kathleen Valley

.... (22)

84

64

168

71

97

60

47

46

10

28

32

78

785

Mt. Sir Samuel

.... (40)

103

90

157

77

111

82

65

57

21

23

38

77

901

.Meekafbarra

.... (33)

137

80

155

93

108

91

64

50

16

18

30

58

909

Nannine ....

.... (47)

107

77

132

68

89

95

64

54

22

19

19

35

781

M'arburton Range

*

697

182

330

92

43

54

31

7

2

61

112

118

1 , .'iSO

* Figures for 1941 only.

Omlow

Cue

Mt. Magnet

Sandstone

Lawlers

Vouannii

Doyle’s Well .

Leonora

Murrin Murrin

Laverton

Morgans

Yiindramindra

Kookynie

Davyhurst

Bulnng ....

Zanthus

Kanowna

Kah/oorlie

M'idgieinooltba

Ualladonia

Rawlmna

Loongana

Eucla ....

(b) Central Zone (Summer •autumn rainfall, Mar.-Junc).

(56)

85

107

170

99

147

155

77

45

4

2

4

15

(47)

95

72

102

70

100

117

00

74

28

21

26

56

(47)

94

91

106

62

104

119

05

02

36

25

29

51

(37)

103

71

119

62

108

95

74

87

23

20

37

1 1

\iT>)

91

73

124

86

92

88

63

54

•>•>

25

30

60

(27)

62

72

107

85

116

92

S6

86

27

33

58

62

104

72

184

78

94

76

50

53

16

34

45

68

(44)

81

78

122

80

93

92

61

66

26

28

44

61

(42)

84

72

137

77

100

94

50

66

24

38

59

75

(4-)

87

81

143

89

98

83

58

54

31

33

68

74

(42)

84

68

139

88

94

75

47

54

28

20

57

54

(41)

80

71

130

63

111

89

55

60

32

33

54

73

81

(40)

86

72

129

76

122

98

76

.SO

33

47

58

(45)

71

94

98

84

119

109

85

SO

34

41

49

66

(40)

67

79

98

97

127

114

103

95

40

63

52

68

(45)

48

71

92

81

109

102

78

96

40

70

62

64

55

77

139

89

107

99

59

86

45

63

79

72

(46)

63

66

107

77

113

105

80

94

43

70

58

70

(46)

57

65

109

86

112

107

83

95

45

70

56

66

(44)

60

76

118

108

124

113

80

104

59

96

86

71

(51)

48

51

99

70

99

93

72

91

68

97

76

67

(25)

27

40

43

51

78

65

58

43

68

76

58

56

42

33

58

64

32

25

57

25

58

36

58

68

(63)

55

67

87

105

123

109

93

92

74

75

66

49

910

soi

004

STO

817

^8«

874

s32

876

m

817

860

958

930
1,012

922

973

946

951

1,U95

931
637
615
995

1 X \’ i i .

Thk Vkuktatiox of Wkstkhn Austiovma.

II. — ICKEMKAN I'HOVINC'E — continued.

(c) Weetern Zori>'.

Station.

Years.

Jan.

Feb.

Mar.

A pi.

May.

June.

July.

Am;.

Sept.

Oct.

Nov.

Dee

Total.

Kiaacoi/ne Junction

.... CM)

109

81

153

50

138

119

102

r.2

13

11

5

22

855

Hamclin Fool

.... (50)

20

50

56

38

125

195

153

76

ar>

14

15

~0

701

•CarnarriOf

.... (4H)

34

03

50

49

153

248

155

80

35

18

7

0

808

Wooranu‘1

.... (43)

44

50

59

46

128

183

135

75

20

17

0

20

780

■Shark Hay

... (48)

31

03

50

49

153

248

155

80

.35

18

7

0

808

Jloolardy

... (51)

00

77

99

53

104

145

105

08

10

0

17

31

700

Murttoo

.... (53)

58

59

99

61

110

155

107

79

22

14

18

34

810

Yuln

... (21)

43

54

79

64

101

145

130

85

20

12

31

27

800

Yaliroo

.... (45)

03

83

106

62

122

179

132

105

45

33

28

41

000

Latham

(H)

08

50

82

166

130

206

191

140

30

31

43

20

1.181

Wuhin

.... (15)

28

00

63

87

151

232

196

184

50

01

35

23

1 . 1 85

•({oodlaml'

... (20)

43

30

101

89

140

208

164

122

51

50

40

51

1,085

KuHa

(0)

32

44

41

84

128

210

183

163

53

70

50

45

1,100

Mollerin

... (13)

38

43

48

85

132

204

155

164

55

82

47

42

1 ,005

Wialki

... (10)

50

04

50

80

106

160

175

182

68

77

30

47

1,100

Honnh* Kock ....

77

43

23

34

88

135

161

127

55

57

81

52

033

Majidiua

(20)

50

00

81

91

133

199

189

149

99

00

40

58

1 .25 1

Hcnfubliin

.... (29)

59

78

95

90

135

191

173

152

80

72

38

51

1,214

Mukinlaidin . .

.... (13)

25

40

59

104

120

149

135

141

62

73

04

52

1,024

Kuminoppin . .

.... (29)

43

07

97

95

130

185

174

143

81

80

30

42

1,173

Nuniiarin

.. . (37)

45

05

87

82

146

185

195

149

80

70

40

47

1,200

Hurraro|)|>in

.... (40)

37

53

80

91

158

182

195

151

98

80

41

45

1,223

■\Valijoo)an

. .. (24)

75

70

93

101

135

192

167

135

73

07

48

.50

1,100

Newcarnie

.... (30)

43

50

95

90

134

190

183

146

02

88

38

02

1,217

AVestonia

.... (20)

39

50

103

94

161

197

189

157

04

88

50

48

1,285

Hullllnch

.... (30)

59

59

97

106

134

150

149

128

08

74

03

02

1,140

Sonthorn Cross

.... (52)

49

57

96

82

138

136

140

116

72

05

52

40

1 ,052

Marvel Hooli ....

.... (20)

52

53

127

101

139

164

175

145

85

03

00

52

1 ,255

Lake V alley ....

... (11)

104

38

137

85

169

148

169

178

100

141

02

27

1,358

Lake Kiii«

.... (12)

82

28

130

08

162

156

178

160

95

147

72

05

1,347

.SaluKtn (Jains ....

.... (:>0)

99

00

1 1 1

99

139

149

133

140

101

117

01

77

1.310

Oooltzardie

.... (49)

59

77

95

92

120

114

86

103

50

70

04

07

1,000

Norseman

.... (45)

55

71

89

96

117

114

96

111

72

02

82

72

1,007

Orasspateh

.... (U>)

118

49

115

105

152

179

161

190

1 m

131

138

03

1,51 1

Kyre

.... (42)

81

47

89

87

161

164

117

125

88

82

08

50

1,108

m.-~s()UTii-\vic.s’r i’Kovin('k.

(a) Emtem Zone.

Station.

Years.

Jan.

Feb.

Mar.

Apl.

May.

June.

July.

Auk.

Sept.

0(!t.

Nov.

Dee,

'rr)tal.

Wubin

.... (15)

28

00

63

87

151

232

196

184

50

<11

35

•23

1.185

flabbin

.... (10)

48

04

85

07

128

195

197

210

98

108

55

22

1,307

Koorda

.... (30)

5,5

53

81

75

121

184

181

141

82

70

30

40

1,125

Wyalkat^eliem ....

.... (32)

03

00

87

87

167

235

227

163

100

81

30

40

1,345

Korieloekinj; ....
Trayninu

.... (20)

40

00

107

91

166

238

218

169

100

83

45

30

1,355

.... (31)

51

03

99

89

135

210

100

153

80

80

41

47

1,200

Kellerherrin

.... (40)

34

47

03

70

166

220

219

174

108

86

43

01

1,320

Doodlukine

.... (31)

30

50

00

85

137

194

192

155

85

93

48

01

1 ,235

Merredin

.... (38)

40

40

02

94

154

203

213

156

104

80

44

57

1,202

liruee Hock

.... (27)

51

71

104

80

167

215

215

169

99

98

40

5*2

1,373

Narembeen

.... (14)

20

33

58

71

146

177

208

180

80

97

53

35

1,(00

JCondinin

.... (24)

41

.50

00

01

178

203

207

171

106

102

.50

42

1,340

Myden

... (13)

53

40

01

75

167

174

205

168

82

111

80

50

1,320

Newde>;ate

.... (10)

73

20

144

78

166

182

212

173

108

132

03

47

1,308

Jariamonj;up ....

.... (47)

55

55

120

128

180

177

205

174

149

144

74

70

1,531

llavensthor|H! ....

.... (40)

70

01

142

120

172

155

180

176

143

156

101

05

1,,580

Oll^reru|l

.... (27)

7S

01

120

1 10

201

197

200

172

150

147

80

48

1,504

Mollerton

42

35

1 1 1

172

134

173

254

251

103

135

41

30

1,520

(h) Centra} Zone.

Ajana

.... (21)

32

37

92

78

213

321

224

176

70

40

24

27

1,220

Yuna

.... (32)

40

50

95

68

190

312

263

181

00

0*2

:io

21

1,421

M'lllewa

... (40)

45

52

78

05

182

267

234

180

91

56

28

20

1.208

NauKettv

.... (38)

40

20

83

80

194

317

270

199

119

08

J1

10

1,455

Canna

.... (22)

52

03

138

97

174

306

242

172

80

00

40

40

1,485

Mlnuenew

.... (40)

32

38

71

81

219

359

318

245

131

81

40

27

1,042

Moraw"

.... (28)

00

73

113

88

172

242

225

167

01

04

57

40

1 ..202

Pereniori

.... (24)

41

72

107

100

173

265

204

167

80

52

40

22

1,242

Three Spriier.s

... (3.5)

54

05

78

92

191

318

292

224

112

74

115

22

1,507

Carnamah

.... (53)

45

54

82

84

206

321

279

229

125

72

40

20

i ,.580

<'oorou' .

.... (30)

45

00

83

95

183

326

283

218

127

82

40

45

1,.587

HnnDne

.... (20)

51

73

112

94

163

276

219

193

85

71

IMi

27

1,407

Watheron

.... (20)

33

5t5

101

92

217

339

294

221

127

85

28

1,020

Dalwallinii

. .. (20)

55

74

00

82

152

276

234

190

104

74

52

27

1,421

Miliny

... (11)

00

70

100

103

191

296

253

257

100

VI

40

44

1..580

Lallidu

.... (20)

50

57

75

91

157

258

224

165

95

77

20

24

1,222

Kondut

(0)

53

50

90

100

186

224

225

222

78

70

40

25

1,284

Wom;an Hil!« ....

. .. (20)

30

50

83

78

189

289

262

208

110

80

40

48

1,470

Cadoux ....

.... (12)

42

51

02

123

165

236

224

208

87

70

51

50

1,281

CalinLuri

... (12)

20

40

83

03

232

329

325

283

119

103

40

27

1,721

Dooinallinir

.... (37)

40

52

70

74

166

276

248

203

106

92

28

48

1,418

Dowerln

.... (38)

41

02

81

78

194

266

260

199

117

80

40

52

1,470

MnlKart

.... (30)

34

05

80

87

246

337

367

273

156

107

44

42

1,844

Ixviii

Charles Austin Gardner,

III.— SOUTH-WEST PROVINCE—

(b) Central Zone — continued.

Station.

Years.

Jan.

Feb.

Mar.

A pi.

May.

June.

J Illy.

Aug.

Sept.

Oct.

Nov.

Dec.

Total..

Minnivale

(30)

52

62

91

82

162

254

237

188

105

79

4-1

43

1.396

Sortham

(61)

34

40

78

84

226

323

342

256

151

100

40

37

1,711

Meckering

(44)

28

47

70

86

200

281

293

223

126

97

44

31

1..526-

Cunderdin

(28)

43

60

87

87

197

245

264

200

95

104

41

35

1,4.58

Tammin

(30)

42

48

90

89

164

230

236

176

106

103

45

49

1 .378

Beverley

(56)

33

42

68

88

230

320

325

250

157

102

49

24

1.688

Quairading

(31)

43

59

89

82

198

236

282

214

117

115

4.3

40

1,518

Pingelly

(51)

33

40

60

00

238

320

331

259

174

107

49

47

1,766

Corrigin

(32)

47

53

95

85

204

244

264

212

129

106

64

66

1,569-

Kulin

(22)

57

63

106

86

233

227

237

174

121

112

62

68

1,546

Wickepin

(30)

41

51

84

103

229

284

295

222

145

103

51

48

1,656

Dudinin

(26)

60

45

90

80

188

265

275

203

135

123

55

37

1,556

Wagin

(51)

36

53

02

125

221

300

303

240

181

126

47

41

1,765-

Diimbleyung ....

(32)

53

55

94

122

206

237

260

215

144

131

61

60

1,638

Kukerin

(20)

67

70

108

105

205

245

253

220

158

131

79

66

1,707

Lake Grace

(27)

60

52

122

97

195

211

210

178

122

112

56

52

1,476

Katanninq

(50)

44

52

103

118

240

297

306

246

189

154

64

62

1,873

Broome Hill ....

(51)

44

57

102

118

221

284

281

233

198

154

70

52

1,814

Nyabing

(20)

56

57

06

108

222

246

262

217

152

130

73

59

1,678

Plngrup

(14)

83

30

1.58

81

204

203

234

199

140

139

77

34

1,.582 .

Tambellup

(31)

72

63

no

126

242

263

267

221

182

157

78

66

1,847 ,

Gnowangerup ....

(20)

68

64

106

137

207

229

222

182

156

142

71

64

1.648 i

Borden

(17)

68

35

115

86

156

182

190

135

127

130

69

73

1.366 ;

Cranbrook

(40)

57

64

106

1.34

238

289

310

256

210

179

92

63

1.998 ,

Hopetoun

(40)

68

62

144

160

238

257

255

227

193

169

103

92

1 .968

Ksperance

(5S)

70

69

115

175

326

407

401

381

271

214

104

86

2,619 :

(0)

Savannah WoodlamI Zone.

Northampton ....

(60)

23

42

55

70

261

453

415

324

184

97

35

22

1,999

Chapman

(36)

28

30

70

70

233

422

388

270

152

91

37

27

1,827

Grraldton

(64)

22

20

57

92

272

477

379

279

128

70

26

15

1,846

Greenough

(60)

16

31

56

91

290

498

391

300

157

80

27

15

1,479

Dongarra

(58)

16

36

58

84

286

463

373

273

152

77

28

16

1.395

Dandarragan ....

(44)

30

41

82

87

283

422

392

336

219

140

54

48

2,134

Moora

(44)

43

44

78

81

239

373

356

265

169

107

41

36

1,832

Barberton

(30)

48

48

77

87

233

355

344

264

158

99

50

40

1 ,803

New Norcia ....

(50)

30

41

82

87

283

422

392

336

219

140

54

48

2,134

'I’oodyay

(62)

31

38

80

05

284

402

429

349

202

141

49

38

2,138

York

(65)

33

46

73

86

239

331

337

280

154

104

4.')

41

1,769

Brookton

(33)

43

43

71

94

249

344

355

267

162

118

48

43

1,837

Wandering

(53)

35

42

03

128

330

461

475

391

283

194

68

60

2,. 560

Narrogin

(50)

38

56

88

112

262

344

371

292

209

141

55

47

2.015

Williams

(57)

29

43

75

122

277

384

397

329

227

143

65

48

1.387

Arthur ....

(51)

32

51

00

116

248

343

344

282

201

143

51

42

1,943

Darkan

(40)

32

38

71

128

301

394

442

343

241

170

58

37

2,255

.Kojonuj)

(57)

48

50

05

132

277

358

358

307

236

185

84

6]

2,200

Kendenuj)

(41)

67

50

123

173

303

330

381

321

282

246

108

83

2,476

Mount Barker

(55)

00

89

145

208

339

386

-417

370

334

290

143

106

2,917

(d) Forest

ay\d

Coastal Districts.

0 ingin ....

(53)

31

42

76

121

433

615

633

500

304

221

57

. 45

3,078

Chidlow

(35)

46

64

85

1.56

495

735

779

636

400

258

89

53

3,796

Mundaring

(36)

36

67

101

204

574

818

895

719

473

307

93

65

4,352

Kalamunda

(33)

47

54

05

209

603

827

838

690

450

302

95

77

4.287

Guildford

(62)

34

40

67

165

485

679

677

559

346

225

60

47

3.303

Perth (Observatory)

(66)

33

40

81

171

505

709

672

571

336

219

76

54

3.467

Fremantle

(67)

29

41

69

1,58

445

636

619

473

277

173

62

45

3.027

Rottnest

(60)

24

30

54

141

439

641

599

443

255

151

57

38

2,881

Armadale

(41)

20

35

74

147

539

738

752

586

380

234

66

44

3,624

Rockingham

(44)

34

20

60

1.53

506

697

685

499

322

194

65

38

3,291

.Tarrahdale

(50)

52

53

102

222

645

896

920

781

540

360

114

72

4,757

Mandurah

(52)

24

36

70

164

524

762

696

536

371

224

69

44

3.529

Binjarra

(63)

34

42

82

186

539

761

731

616

386

253

90

58

3,778

DwelHngup

(15)

47

62

105

300

720

1,024

1.082

799

586

324

128

47

5.180

Marradong

(44)

36

44

60

142

409

549

590

457

323

203

70

47

2,939

Brunswick

(33)

46

64

85

181

581

810

802

625

440

290

no

50

4.084

Bunbury

(65)

40

.50

03

170

512

706

673

514

348

232

94

54

3,486 j

Collie

(42)

57

58

06

102

522

701

755

584

460

308

114

62

3.909

Cape Naturaliste

(38)

41

60

03

140

479

682

673

446

301

217

83

48

3.272

Busselton

(61)

42

45

85

145

470

674

649

459

301

226

85

48

3,229

Donnybrook

(41)

45

63

111

176

569

768

776

608

463

298

116

61

4,054

Greenbusbes

(40)

59

70

112

195

519

681

675

588

441

319

119

77

3,855

Cape Leevu'ui ....

(45)

64

77

116

218

582

715

729

541

341

283

121

79

3.866

Manjininp

(26)

82

83

no

221

588

711

710

633

479

347

164

88

4.225

Pemberton

(6)

160

57

177

204

699

787

810

888

493

422

219

105

5.111

Bridgetown

(54)

58

50

107

170

458

591

581

514

384

291

no

74

.3.397

Bovup Brook ....

(28)

50

58

87

148

366

484

491

407

283

212

88

34

2,708

Denmark ....

(40)

126

135

217

323

570

673

728

646

504

429

181

140

4.672

Albany ....

(65)

101

88

159

275

502

543

559

530

408

324

146

115

3.750

Kcli))se Island

(52)

87

80

144

239

437

474

527

469

351

292

125

107

3.332

riTK Vegetation of Western Australia,

Lxix.

TABLE II.—ANNUAL AND SEASONAL RAINFALL, AND ANNUAL RAINFALL

VARIATION.

The rainfall is given in points (100 points to an inch) and mm. The wet season indicates
the four consecutive wettest months, and the dry season the four consective driest months. The
annual rainfall variation per cent, departure from the mean is given within close limits, (after
GeniiUi). The column marked “ % ” indicates the proportion of the rainfall for the four wettest
months as a fraction of the annual rainfall.

I.—NORTHERN province.

—

Annual.

Wet Season.

Per

cent.

Dry Season.

Ann.

var.

Port Oeortre IV

])oints.

5,039

mm.

1,280

Dec. -Mar.

points.
.... 4,391

lum.

1,115

87

July-Oct.

point.s.

79

mm.

20

15-20

Vlount Barnett . ..

3,457

879

.... 2,761

701

79

June-Sept.

57

14

15-20

Kimberley Downs ....

2,500

637

.... 2,124

540

85

,Iuly-Oct.

40

10

20-25

Beasle Bay

2,778

706

.... 2,243

570

81

83

21

25-30

Turkey Creek

2,714

690

.... 2,242

570

82

June-Sept.

81

20

20-25

yndham*

2,510

637

.... 2,229

566

85

43

11

15-20

Derby*

2,530

643

.... 2,141

544

85

•July-Oct.

43

11

25-30

Broome*

2.301

.584

.... 1,934

491

87

39

10

30-35

Fitzroy Crossina ....

2,110

536

.... 1,827

464

87

June-Sept.

62

16

20-25

Hall’s Creek*

1,961

499

.... 1,591

404

81

75

19

25-30

La Grange

1,870

475

.... 1.517

385

92

July-Oct.

47

12

30-35

Anna Plains

1,447

368

.... 1,096

264

75

45

11

30-35

Wollal

1,206

306

978

242

81

Aug.-Nov.

26

7

Marble Bar*

1,315

334

918

233

69

78

20

30-35

Nnllagine*

1,297

329

909

231

70

July-Oct.

90

23

30-35

Hamersiev ....

1,337

340

983

250

74

93

24

30-35

Condon

1,191

303

•Tan.-Apl.

879

223

75

Aug.-Nov.

32

8

35-40

Port Hedland*

1,193

303

.... 827

210

69

Sept. -Dec.

37

12

3.5-40

Roebonrne*

1,168

297

831

219

71

Aug.-Nov.

42

11

40-45

Bamboo Creek

1,420

361

947

241

67

95

24

30-35

Tamb ey

1,562

397

. .. 1,071

272

69

89

23

25-35

Mll.stream

1,477

375

.... 1,001

254

67

74

19

2.5-35

'Whim Creek

1,467

373

.... 1,093

278

74

66

17

40-50

Red Hill

1.356

344

909

231

67

65

17

45-50

Bonny Downs

1,131

287

Dec. -Mar.

.... 772

196

68

Jiily-bct.

78

20

35-40

Miindiwindi*

points.

1,021

mm.

2.59

11.— EREMEAN PROVINC
(a) Northern Zone.

I>oints. mm.
Dec. -Mar. .... 632 161

E.

62

July-Oct.

points, mm.
122 31

35-40

Ethel Creek

1,001

254

644

163

64

82

21

35-40

Roy Hill

1,001

2.54

667

169

67

Aug.-Nov.

82

21

35-40

Three Rivers

872

221

482

122

55

82

21

35-40

-Kathleen Valiev ....

785

199

394

101

50

116

29

3.5-40

Mardie

870

220

.Jan. -April

605

154

58

Sept. -Dec.

40

10

35-40

V inning Pool*

986

205

512

130

51

56

14

40-45

^ laming Head

1,215

309

752

191

61

10

2

30-35

V andagee . ..

901

228

436

111

48

61

15

40-45

Peak Hill

940

239

498

126

53

Aug.-Nov.

106

27

35-40

^\il^na*

938

238

514

131

55

111

28

35-40

Meekatliarra*

909

231

465

118

51

Sejit.-Dec.

122

31

35-40

Nannine

781

198

384

98

49

95

24

35-40

Mt. Sir Samuel

901

228

.... 427

108

47

Aug.-Nov.

139

35

35-40

Onslow*

points,

910

mm.

230

(b) Central Zone.

points, mm.
Mar.-.June .... 571 145

63

Sept. -Dec.

point.s.
.... 25

mm.

6

55-60

<’ue*

851

216

389

99

46

131

33

3,5-40

Mount Magnet

804

230

391

99

43

141

36

30-35

Sandstone . ..

879

223

* >

384

98

44

no

27

25-30

Lawlers

817

208

390

99

47

Aug.-Nov.

140

36

2.5-30

^ ouanmi

886

225

400

102

35

Sept.-Dee.

180

46

2.5-30

Doyles ^\ell

874

222

f )

432

no

50

Aug.-Nov.

148

38

25-30

Leonora

832

211

> •

387

98

47

Sept.-Dee.

159

40

30-35

Malcolm

876

222

408

104

46

30-35

Murrin Murrin

876

222

• 1

408

104

46

Aug.-Nov.

186

47

30-35

Laverton . .

899

228

413

105

46

July-Oct,

176

45

30-35

Morgans

817

208

396

101

49

Aug.-Nov.

168

43

30-35

\ undramindra

860

218

393

100

45

188

48

30-35

Kookynie ...

958

243

425

108

44

218

30-35

Menzies*

930

236

410

102

43

Sept.-Dee.

190

48

25-30

Davyhurst

1,012

257

« 1

436

111

43

232

59

20-25

Bulong

922

234

384

98

42

Oct. -Jan.

244

62

20-25

Zanthus

973

247

s *

434

no

45

Sept. -Dec.

259

66

25-30

Kanowna ...

946

240

402

102

43

241

61

2.5-30

Kalgoorlie*

951

242

414

105

43

237

60

20-25

idgiemooltha

1,095

278

463

117

54

Nov. -Feb.

293

74

20-25

Balladonia*

931

236

.... 361

92

39

242

61

20-25

Rawlinna*

L<xuigana ....

637

162

.... 262

66

41

186

47

25-30

615

1.56

.... 240

61

39

Sept. -Dec.

154

39

30-35

Kucla

995

253

April-.! Illy

430

109

44

Nov. -Feb.

237

60

20-25

Ixx.

Charles Austin Gardner.

1I.~EREMEAN PROVLNC'E—

(c) Western Zone,

—

Annual.

Wet Season.

Per.

cent.

Dry

Season.

Ann.

var.

Gascovne Junction

points.

855

mm.

217

points.

May-Aug 411

mm.

104

48

Sept.-Dee.

points.

51

mm.

13

35-40

Hameiin Pool*

791

201

„ .... 548

139

69

Oct. -Jan.

.... 94

24

30-35

Carnarvon*

898

228

636

162

71

95

24

35-40

Wooramel ....

789

200

M .... 521

132

66

Sept.-Dee.

69

17

35-40

Shark Bay

898

228

„ .... 636

162

71

Oct.-Jan.

65

16

35-40

Boolardy ....

790

201

422

107

54

Sept.-Dee.

.... 73

18

35-40

Murgoo

Yuin

810

207

„ .... 451

114

63

.... 88

22

35-40

800

203

461

117

58

’ 1

.... 99

25

25-30

Yalgoo

990

254

.... 538

136

53

.... 147

37

25-30

Wubin

1,185

301

763

194

64

Oct.-Jan.

.... 147

37

20-25-

Goodlands ....

1,085

276

,, .... 624

158

;)/

Nov.-Feb.

.... 170

43

25-30

Kulja*

1,109

282

„ .... 684

174

62

Dec. -Mar.

.... 162

41

25-30

Mollerin

1,095

278

„ .... 655

166

59

Nov.-Feb.

.... 170

43

25-30

Wialki

1.109

282

„ .... 623

158

57

.... 200

51

25-30

Bonnie Rock

938

237

„ .... 511

130

54

Jan.-April

.... 177

45

25-30

Gabbin

1,307

332

„ .... 730

185

59

Nov.-Feb.

.... 190

48

Koorda

1.125

286

., .... 627

159

11

.... 190

48

20-25

Mandiga

1,251

318

,, .... 670

170

.s6

>1

.... 220

56

25-30

Bencubbin ....

1,2U

308

651

165

53

Oct.-Jan.

220

56

25-30

Mukinbudin

1,024

260

545

138

53

Dec. -Mar.

.... 176

45

25-30

Kuniinoppin

1,173

298

.... 632

160

54

Nov.-Feb.

.... 188

48

20-25

Niingarin ....

1,209

307

,, .... 675

171

56

11

.... 197

50

20-25-

Burracoppin

1,223

311

., .... 686

174

56

11

.... 176

45

20-25

Walgoolan ....

1,1 90

304

619

157

52

Sept. -Dec.

.... 238

60

20-25

Newcamie ....

1.217

309

,, .... 653

166

53

Nov.-Feb.

.... 199

51

20-25

Westonia ....

1,285

327

,, .... 704

179

55

11

.... 203

51

20-25

Bullfinch ....

1.141

299

„ .... 561

142

49

1 1

.... 243

62

20-25

Southern Cross*

1.052

267

530

134

50

1 1

.... 207

53

20-25

Marvel Loch

1,255

319

623

158

51

>1

.... 226

57

20-25

Lake King

1,347

342

,, .... 654

166

48

11

.... 247

63

15-20

Lake Varley

1,358

345

664

168

49

>1

.... 231

59

1 .5-20

Salmon Gums

1,316

334

„ .... 627

159

49

)1

.... 327

83

15-20

Cftolgardie ....

1,009

256

„ .... 423

107

42

1 1

.... 267

68

] 5-20

Norseman

1,067

271

„ .... 438

111

41

.... 280

71

20-25

Grasspatch

1,511

384

682

173

45

>1

.... 368

93

20-25

Eyre

1,168

297

„ .... 567

144

41

>1

.... 255

65

20-25

Latham

Wu)>in

Galibin

('adonx

Wyalkatchem

Korrelockinp:

Trayninfj ....

Kellerberrin*

Doodlakine

JVIerredin ....

Druce Rock

Narembeen

Kondinin ....

Hyden

Newdegate

Jarramongup

Biintine

Ongcnip

in.-SOUTH-WKSTERN PROVINCE.

(a) Easteryi Zone.

points.

mm.

points.

mm.

Sept. -Dec.

points.

mm.

1,181

300

31ay-Aug.

667

169

57

.... 142

36

1,185

301

763

194

65

Oct.-Jan.

.... 147

37

1,307

332

730

185

59

Nov.-Feb.

.... 190

48

1,381

351

.... 833

212

60

> 1

.... 206

52

1 ,345

342

725

184

54

11

.... 208

53

1,355

344 .

781

198

56

1 <

.... 193

49

1,266

321

698

177

55

.... 202

51

1,329

338

779

198

58

.... 185

47

1,235

314

678

172

55

.... 204

52

1,292

328

726

184

59

.... 187

47

1,373

349

766

195

56

13

.... 220

56

1.166

296

710

179

.63

1 1

.... 150

38

1,340

340

750

193

56

1 '

.... 192

49

1,320

335

) 1

.... 714

181

46

>1

.... 247

63

1,398

355

733

186

52

1 1

.... 243

62

1,531

389

736

187

48

1 1

.... 254

65

1,407

357

770

196

49

Oct.-Jan.

.... 192

49

1,564

397

11

.... 770

106

49

Nov.-Feb.

.... 267

68

20-25-
20-25
20-25
20-25
20-25
20-25
20-25
20-25
20-25
15-20
15-20
15-20
15-20
1 5-20
1 5-20
20-25
1 5-20

(•))

points.

mm.

Ajana

1.336

339

May-Aug

Ymia

1,421

361

Mullewa

1,308

332

»1

Canna

1,485

377

' >

Mingenew*

1,642

417

Morawa*

1,392

353

11

Pereni'ori ....

1,343

341

1 1

Three Springs

1 ,567

398

? »

(’arnaipah*

('oorow

1,580

1,567

401

398

* 1
1 )

Watheroo

1,636

415

11

Dalwallinu

1.421

361

J 3

liallidu

1,322

336

>1

Miling

1,586

403

11

Kondut

1,384

351

i>

Walebing ....

2,013

511

n

Wongan Hills

1,476

375

1 1

Cadoux

1,381

352

'1

Calingiri

1,731

440

Central Zone.

])oints.

mm.

points.

mm.

20-25

933

237

70

Nov

.-Feb. ...

. 120

30

946

240

67

M

160

41

20-25

863

219

66

155

39

20-25

894

227

60

Oct.

-Jan. ...

. 207

52

20-25

1,141

290

69

Nov

.-Feb. ...

. 137

35

20-25

806

205

58

Oct.

-Jan ...

. 221

56

20-25

809

205

60

. 172

44

20-25

1 ,025

260

66

Nox

-Feb. ...

106

50

20-25

1 ,035

263

65

181

46

20-25

1,010

255

65

11

. 190

48

20-25

1,071

272

65

51

. 160

41

20-25

852

216

60

Oct.

-Jan. ..

. 219

56

20-25

804

204

61

Nov

.-Feb. ..

. 180

46

20-25

996

253

63

J 1

. 214

54

20-25

857

218

62

11 • •

. 183

46

20-25

1,324

336

65

11

. 188

48

15-20

948

241

64

>1

. 180

46

20-25

833

207

63

11

. 206

52

20-25

1,169

297

67

11 * •

. 164

42

15-20

The Vegetatiox of Western Australia.

Ixxi

III.— north-western province— coniinwcrf.

(b) Central Zone — continued.

—

Annual.

Wet Season.

Per

cent.

Dry Season.

Ann.

var.

points.

mm.

points.

ram.

points, mm.

Goooialling

1,418

360

Mav'Aiig.

893

227

63

Nov.-Feb

178

45

20-25

Dowerin

1,479

376

.... 919

233

60

204

52

20-25

Northam

1,711

435

.... 1,147

291

67

151

38

15-20'

Meckering

1,526

388

.... 997

253

65

150

38

20-25-

Cunderdin ....

1.458

370

.... 906

230

62

179

45

20-25

Tammin

1,378

350

806

205

58

184

47

20-25

BeverJev

1,688

429

.... 1127

286

67

148

38

15-20

Quairading

1,518

385

930

236

62

185

47

15-20

Pingelly

1,766

448

.... 1,148

292

65

169

43

1.5-20

Corrigin

1,569

398

924

235

59

230

58

15-20

Kulin

1,546

393

871

221

56

250

63

15-20

Wickepin ....

1,656

421

.... 1,020

262

62

191

49

15-20

Diidinin

1,556

395

.... 931

236

59

197

50

10-15

Wagin

1,765

448

.... 1,064

270

60

177

45

15-20

Diimbleyung

1,638

416

918

233

56

229

58

15-20

Kukerin

1,707

433

.... 933

237

54

282

72

15-20

Lake Grace

1,476

375

794

202

54

229

58

15-20

Katanning*

1,873

476

.... 1,089

277

58

222

56

15-20

Broome Hill

1,814

461

.... 1,019

259

56

223

56

15-20'

Nyabing

1,678

426

947

240

56

245

62

10-15

Pingrup

1,582

402

.... 840

213

52

224

57

10-15

Tambellnp ....

1,847

469

993

252

54

279

71

10-15.

Gnowangerup

1,648

418

840

213

57

267

68

15-20

Borden

1,366

337

663

168

57

245

62

15-20.

Cranbrook

1,998

.507

.... 1,093

277

55

276

70

10-15

Hopetonn

1,968

500

977

248

50

325

82

10-15

Esperance*

2,619

665

.... 1,515

385

58

329

83

10-15

Northampton

1,990

505

.... 1,453

369

73

122

31

15-20'

Chapman (R.S.) ....

1,827

464

.... 1,313

335

72

131

33

15-20'

Geraldton*

1,846

460

... 1,407

357

76

92

23

15-20'

Greenongh ....

1,952

496

.... 1,479

376

75

89

22

20-25

Dongarra ....

1,862

473

.... 1,395

354

75

96

24

20-25

Dandarragan

2,134

542

.... 1,433

364

67

173

44

15-20'

Moora

1,832

465

.... 1,233

313

67

164

41

15-20

Barberton

1,803

458

.... 1,196

304

66

186

47

15-20-

New Norcia

2,134

542

.... 1,433

364

56

173

44

1 5-20

Toodvay

2,138

543

.... 1,464

372

68

156

40

15-20

York

1,760

449

.... 1,187

301

67

165

42

15-20

Brookton ....

1,837

467

.... 1,215

309

66

177

45

15-20.

Wandering

2,560

650

.... 1,657

421

64

205

52

1 .5-20

Narrogin

2,015

512

.... 1,269

322

63

196

50

10-15

Williams

2,139

543

.... 1,387

254

65

185

47

15-20

Arthur

1,943

493

.... 1,217

309

62

176

45

15-20

Darkan

2,255

573

.... 1,480

276

65

165

42

15-20

Koionup

2,200

559

.... 1,300

330

59

252

64

15-20

Kendeniip

2,476

629

.... 1,335

339

54

317

81

10-15-

Mount Barker

2,917

741

-

.... 1,512

384

52

....

428

109

10-15.

(0)

Western Zone.

points, mm.

points.

mm.

points, ram.

Gingin

3,078

782

Mav-Aug.

.... 2,181

554

71

Nov .-Feb

175

44

15-20

Chidlow

3.796

964

.... 2,645

672

70

252

64

15-20

Mundaring

4,352

1,105

.... 3,006

763

69

261

66

1 5-20'

Kalamunda

4,287

1,080

.... 2,958

751

69

273

69

1 5-20

Guildford

3,393

862

.... 2,400

610

71

190

48

1 5-20

Perth Observatory

3,467

881

.... 2,457

624

71

203

51

15-20'

Fremantle

3,027

769

.... 2,173

552

72

177

45

1 5-20

Rottnest

2,881

732

.... 2,122

539

75

158

40

1

Armadale

3,624

932

.... 2,615

664

71 .

174

44

1.5-20

Rockingham

3,291

836

.... 2,387

606

73

166

42

15-20

Jarrahdale

4,757

1,208

.... 3,242

823

68

Dec. -Mar

369

94

10-1.5

Mandurah ....

3,529

896

.... 2.518

639

69

Nov.-Feb

173

44

10-1.5

Pinjarra

3,778

959

.... 2,647

672

71

Dec. -Mar

216

55

10-15

Dwellingup

5,180

1,316

.... 3,625

921

71

261

66

10-15-

Marradong

2,939

747

.... 2,005

509

68

196

49

10-1.5-

Brunswick ....

4,084

1,037

.... 2,818

716

69

254

62

10-15.

Bunbury ....

3,486

885

.... 2,405

611

70

237

60

10-1 5

Collie

3,909

993

.... 2,562

651

65

273

69

10-15

Cape Naturaliste ....

3,272

831

.... 2,280

579

70

242

61

10-15

Busselton

3,229

820

.... 2,252

572

69

220

56

10-15

Donnybrook

4,054

1,030

.... 2,721

691

67

280

71

10-15

Greenbushes

3,855

979

.... 2,463

626

64

318

81

10-15

Cape Leeuwin

3,866

982

.... 2,567

652

64

336

85

10-15

Manjimup ....

4,225

1,073

.... 2,642

671

63

372

95

10-15

Pemberton

5,111

1,298

.... 3,184

809

45

499

127

10-1 5

Bridgetown

3,397

863

.... 2,144

545

64

298

76

10-15

Boyup Brook

2,708

688

.... 1,748

444

64

229

58

10-15

Denmark

4,672

1,187

.... 2,617

665

47

Nov.-Feb. ..

582

148

10-15

Albany

3,750

952

.... 2,134

542

57

450

114

10-15

Eclipse Island

3,332

846

5 >

.... 1,907

484

58

”

399

101

10-15

I

Ixxii.

CirAHLKs Austix Gardner.

TABLE III.— RELATIVE HUMIDITV.

(«) Readings taken at 9-0 a.m.

{b) Readings taken at 3-0 p.m.

The higher readings are in heavy type.

(Those station.^ with less than three years records are marked t. The figures for 1941 alone
are given.)

•Station.

Jan.

l-’eb.

Mar.

Apl.

May.

.Tune.

July.

Aug.

Sept.

Oft.

Nov.

Dee.

Mean,

"Wymlham

(a)

54

69

56

46

41

42

40

43

44

51

56

55

.52

(b)

54

54

40

38

37

37

35

30

43

47

50

52

15

Hall’.s Creek

(n)

51

52

46

35

37

40

37

33

29

32

34

45

42

(b)

30

33

27

29

32

30

20

24

26

22

25

28

Derby

(a)

70

79

66

52

47

47

47

46

47

51

56

63

55

(b)

58

50

53

42

40

40

40

38

30

43

50

54

46

liroome

(a)

72

73

69

54

52

54

52

53

52

57

60

66

f)9

(b)

07

07

50

45

44

40

43

42

44

52

57

02

52

Anna Plains

(a)

00

00

01

43

42

48

40

42

30

45

49

54

51

Port Hedlami . ..

(a)

02

02

50

40

47

50

47

47

44

40

48

50

51

(b)

63

63

50

49

49

47

47

49

49

53

55

61

53

Marble Bar

(a)

44

46

43

39

46

48

45

39

32

30

29

36

40

(b)

21

28

27

20

31

33

30

20

22

20

10

23

26

Roeboiirm*

(a)

49

51

52

43

46

50

45

45

37

37

36

42

44

(b)

41

42

43

30

38

40

30

30

20

33

33

37

37

Mimdiwimli

(a)

29

30

34

33

42

49

44

38

26

22

20

25

44

(b)

17

18

20

19

25

28

23

10

13

12

11

15

18

■Onslow

(a)

52

54

54

52

56

60

57

52

46

44

44

47

52

(b)

52

54

53

50

52

54

51

47

45

44

48

50

50

Winning Pool

(a)

45

49

49

45

50

54

48

43

41

38

37

40

45

(b)

24

20

28

27

33

37

32

30

25

23

20

22

28

<’arnarvon

(a)

00

58

57

00

66

66

62

57

55

50

57

59

(b)

61

62

61

59

00

50

57

50

59

59

60

62

60

•Oascoyne Jnnet.t

(a)

34

48

36

51

60

61

59

45

46

43

40

39

55

(b)

10

21

23

32

45

40

40

20

20

30

20

20

31

Hamelin Pool ....

(a)

43

46

48

52

62

69

60

64

55

49

44

43

54

(b)

.30

38

40

45

54

00

50

54

47

43

30

38

46

Meekatharra

(a)

31

29

58

41

50

60

61

53

40

33

31

27

41

(b)

18

10

33

27

34

41

43

35

23

20

10

17

27

Wiluna ....

(a)

35

37

42

46

53

59

57

51

42

38

34

33

44

(b)

25

27

30

33

30

42

30

35

31

28

25

24

31

baverton

(a)

36

37

44

47

54

61

60

52

41

37

35

34

4.)

(b)

24

25

30

33

40

40

43

37

28

20

24

24

32

■Cue

(a)

36

39

43

49

59

68

67

60

49

43

37

35

49

(b)

25

20

20

34

42

40

48

42

33

20

25

24

34

Menzies ....

(a)

37

39

45

52

59

66

65

57

45

41

36

36

48

(b)

23

24

20

35

42

48

45

30

30

20

23

22

32

Yalgoo

(a)

40

44

51

56

66

76

77

71

58

49

42

39

56

(b)

25

28

33

37

44

54

54

47

38

31

26

24

37

Rawlitma

(a)

49

52

59

59

64

70

70

63

51

48

47

48

57

(b)

23

20

35

31

35

44

40

35

20

21

21

30

Euela

(a)

50

01

00

61

65

71

70

63

56

53

55

50

61

(b)

62

63

62

58

50

57

50

54

54

57

60

61

58

Balladonia

(a)

54

58

64

69

74

77

77

70

60

55

53

53

64

(b)

34

30

42

40

52

58

y;>

48

40

37

35

35

43

Kalgoorlie,

(a)

46

49

55

59

66

74

74

66

53

48

44

43

.56

(b)

27

30

34

30

40

52

50

44

34

31

28

27

Ol

;South(‘rn Cross . ..

fa)

46

51

56

61

73

81

83

76

63

55

49

46

62

(b)

20

32

40

42

51

60

00

54

43

38

34

30

Knljaf ...

(a)

37

46

47

65

60

86

87

71

61

47

42

41

57

(b)

10

24

25

40

45

04

52

47

44

26

23

36

Salmon Gumsf ....

(a)

05

74

81

70

80

07

03

50

47

51

65

The Vegetation of Western Australia,

Ixxiii

TABLE III.— RELATIVE continued.

Station.

Jan.

Feb.

Mar.

Apl.

May.

June.

July.

Aug.

Sept.

Oct.

Nov.

Dec.

Moan.

Kellerberrin

(a)

51

54

62

65

78

83

86

80

70

58

49

48

65

(b)

30

32

38

42

54

61

62

57

47

38

32

34

44

Northam

(a)

65

71

58

66

79

85

87

80

71

60

50

49

76

(b)

36

42

31

45

57

65

65

59

53

45

35

33

47

Geraldton

(a)

54

56

57

58

65

70

70

68

63

58

55

55

61

(b)

60

60

60

58

58

6f)

60

59

59

59

60

60

59

Watheroo

(a)

38

40

48

53

68

77

80

71

60

50

40

37

5S

Perth Observatory

(a)

51

53

57

62

71

76

77

72

67

61

54

51

63

(b)

44

42

45

49

58

63

63

61

58

55

49

47

53

Carnamaht

fa)

33

47

45

67

62

85

81

78

67

50

46

44

59'

(b)

16

20

21

39

45

66

59

53

45

33

31

24

38

Katanning

(a)

55

59

65

72

81

86

87

82

76

67

58

55

70-

(b)

35

37

43

50

62

69

69

65

58

52

40

37

51

Biinbury

(a)

50

61

64

72

77

80

80

77

73

68

63

61

70*

(b)

57

56

58

63

68

70

71

69

67

65

60

59

64

Esperance

(a)

62

63

66

70

75

78

78

74

69

65

62

61

69

<b)

63

64

64

65

64

65

65

62

63

64

64

63

64

CoUief

(a)

69

79

89

88

88

92

88

80

62

57

51

75-

Manjimupt

(a)

63

57

77

77

78

87

90

87

82

74

75

66

75-

Cape Leeuyint . ...

fa)

89

88

88

73

76

86

83

80

85

80

89

89

84

(b)

88

85

83

73

73

82

80

78

77

SO

89

89

81

Albanj*^

(a)

71

72

74

79

81

77

81

79

77

74

71

69

76

(b)

65

66

67

68

69

70

70

68

69

68

67

65

68

TABLE IV.— SATURATION DEFICIT.

(Inches of mercury).

Taken from 9.0 a.ni. and 3.0 and dry bulb readings.

Station.

Jan.

Feb.

Mar.

Apl.

May.

June.

July.

Aug.

Sept.

Oct.

Nov.

Dec.

Mean.

■\Vyndham

. fa)

•417

•380

•446

•663

•629

•538

•525

•569

• 656

• 650

•608

■514

•55a

(b)

•710

•700

•790

•960

•880

•750

•770

•760

•800

•810

•820

•760

•792

Derby

.. fa)

•366

•355

•415

• 566

•539

•448

•438

• 500

•577

•622

•578

•488

•492

(b)

•590

•640

•700

•880

•790

■680

•680

•760

-870

•880

•750

•700

• 78&

Broome ....

.. fa)

•336

•325

•370

•515

•440

•347

■346

•372

•462

•499

•499

•425

•437

(b)

•420

•420

•580

•800

•680

•570

•560

•630

•670

•580

• 550

•520

■591

Hall’s Creek

- fa)

•634

•598

•695

•716

•572

■428

•433

•580

•795

•928

•944

•707

(b)

1-050

1-050

1-050

1-800

•880

•710

•710

•890

1-140

1-300

1-330

1 • 150

1-015-

Anna Plains

- (a)
(b)

■376

■365

•476

•598

■469

-335

■310

■418

•507

■611

•650

■619

•601

Marble Bar

■ ■ fa)

•828

•765

•764

•702

•457

•353

•359

•451

•671

•865

1-085

•977

•690'

fb)

1-600

1-520

1-340

1-210

•870

•700

•690

•870

1-170

1-450

1-690

1-680

1-184

Port Hedland

.. fa)

•497

•516

•574

•621

•465

• 358

•349

•406

• 536

•599

• 687

•603

•518

(b)

•530

•530

•690

•690

•570

•510

•460

•480

• 560

• 550

•600

• 570

-565

Roebour)ie

- (a)

•807

•758

•666

•707

•482

361

•375

•430

•624

•741

•942

•894

•649-

(b)

1-070

1-070

•980

•960

•730

•580

• 600

•680

•920

•980

M80

1160

■885-

Mundiwinii

. fa)

•976

•892

•739

• 574

•366

•2.54

•266

• 332

•543

•708

•964

•995

•634

(b)

1-480

1-450

1 -230

•970

•680

•510

•540

•640

•960

1-150

1-460

1-560

1-005-

Ixxiv

CirAKLFS AI’STIX (iARDXER.

TABLE IV.— SATURATION DEFICIT—

Station.

Jan.

Feb.

Mar.

Apl.

May

June.

July.

Aug.

Sept.

Oct.

Xov.

Dec.

Mean,

■Onslow

fa)

•fi24

■ 576

•557

•516

•365

•273

•267

•341

•449

• 555

•647

•666

•486

(b)

•690

•730

•670

•630

• 550

•420

•410

• 500

•570

•660

•670

•700

•582

ATiluna

fa)

♦826

•769

■615

•428

•269

•179

•180

•235

•36.3

•.501

•719

•823

•492

(b)

l-.SOO

1*270

1 -040

790

• 500

•380

•370

•460

•670

•870

1-110

1-310

•785

Meekatharra

fa)

•879

■863

•561

•4.50

•280

•170

•151

•206

•34f>

•495

■677

•846

•493

(b)

1-410

1^480

1-080

•830

•570

-.380

•340

•460

•700

•020

1 • 1 60

1-420

•839

'Cue

fa)

•79]

•716

•566

■390

•225

•134

•133

•184

•300

•413

•610

•757

•m

(b)

1-370

1-300

1-040

• 750

• 500

•320

•310

•420

•610

•790

1060

1-310

•755

Laverton

. fa)

•711

•6;52

•496

•383

•2;58

•160

•163

•227

• 357

•461

■621

•709

•434

(b)

1*220

1160

•920

■710

•440

•340

•330

•410

•640

•820

•990

1-200

•719

Alenzies

fa)

•648

•571

•456

•322

•206

• 1 35

•134

•185

-.303

•390

•555

•637

•379

(b)

1-170

1-110

•900

•600

■400

-.300

•290

•390

• 580

•740

l-OlO

1-150

•675

Rawiinna

(a)

•38;")

-.365

•280

■2.38

•168

•113

•103

•J42

•235

•297

•376

•366

•256

(b)

•920

•940

•670

■620

•460

•340

•360

•410

• 550

•740

•900

•990

•640

'Carnarvon

. fa)

•45.5

•427

•435

■397

•291

•199

•185

•1.52

•302

•359

•395

•422

•341

(b)

•420

■430

•460

■430

•350

•320

•310

•280

•330

-.3.50

•370

•390

•363

A’algoo

. fa)

•677

•576

•434

•312

•174

•099

•093

•124

•215

•338

•509

•643

•349

(b)

1-270

1-210

•940

•700

•460

•290

•280

•350

•530

•690

1-010

1-220

•691

'Kalgooriie

. (a)

•492

•434

•332

•249

•163

•099

•095

•1.38

•237

•337

•441

•504

•293

(b)

1-080

1-020

•770

• 570

■390

•270

•270

•340

•510 •

•670

•710

1000

•608

BalladmYja

. (a)

•3;59

■320

•246

•184

•122

■085

•085

•123

•200

•269

•327

•351

■195

(b)

•820

•780

•610

■480

-.3.30

•240

•240

•310

•430

•540

•660

•750

Southern Cross

. (a)

•511

•429

•332

•233

•123

•070

•060

•096

•181

•271

•404

•498

•267

(b)

1-100

l-OOO

•780

•570

■330

•210

•210

•280

•400

• 550

•800

1-020

•551

AVatheroo

. fa)

•588

•544

■419

■307

•159

•092

•080

■120

•199

■297

•4.58

• 560

■319

(b)

Kellerberrin

•428

■378

•275

•198

•099

•061

•051

•076

•136

•233

•369

■428

•226

(b)

1-000

■910

•730

• 540

•300

•210

•210

•240

•370

• 500

•790

•940

•518

Northam

. fa.)
fb)

■4.59

•400

■314

■210

-098

■059

•049

•081

■138

■229

-370

•450

•238

R.speranee

■ (a)

•285

■272

•237

•183

• 1 55

• 095

•092

•114

•157

•199

•250

•281

•193

(b)

•300

•300

•260

•240

•230

•190

•180

•210

•220

•230

•240

•290

•235

*fleraldton

. (a)

• 4.52

•434

•398

•331

•227

•166

• 1 52

•173

•225

•293

•364

•414

•302

fb)

•390

•420

•420

•400

•330

•270

•260

■270

•280

•320

• 340

•360

•330

JCatanning

. (a)

•324

•283

•221

•149

•080

•052

•045

•065

•099

•1.59

•255

•315

•171

(b)

•770

■720

• 550

-.390

•220

• 1 50

•140

•170

•240

•330

• 520

•690

•378

'Pertii Observatory (a)

•417

•392

■323

•244

•144

•101

•096

•120

•167

•224

•320

•398

•246

•590

-630

■ 520

•440

•270

•2^0

•180

•190

•250

•300

•400

•510

•366

Rnnbiiry

. (a)
(b)

•333

■310

■267

•170

•112

•088

•083

•098

•135

•181

•2.58

•309

•195

•420

•420

•390

■290

•200

•160

• 1 50

• 1 50

•190

•230

•290

•390

Bwellingup

- fa)
(b)

■275

•261

■187

■120

•0;')9

•036

-027

•062

■091

■138

■237

-280

■148

'Cape Leenwin

- fa)
•fb)

•089

•093

•077

•099

•087

•070

•081

•089

■074

•085

•104

•090

•094

•190

•190

•180

•140

•130

•090

•090

•120

•120

•120

•170

•180

•140

Albany ....

■ (a)
(b)

•191

•184

■153

•112

•099

•080

•072

•065

•098

•131

•174

•183

■128

•250

•250

•210

•200

■190

•150

•150

•150

•1.50

•190

•200

•240

Based on

Mean 9

•0 a.m.

readings for 1941-1942.

'^-ascoyneJnnrtion

■680

•568

•548

■441

•218

•159

■152

•232

■321

■351

• 584

■691

-421

Kiilja . ..

■ 503

■428

■284

■197

■169

•070

•058

■125

•153

•280

■413

■501

■412

’Carnamah

•567

■438

■254

■205

•160

•062

■073

■088

■154

•237

■419

•525

■270

Salmon Oums

•361

•135

■185

■151

■102

■067

■080

■134

■178

■246

■372

■367

■207

■Collie

■267

•190

■096

■078

■044

•047

■024

■050

■074

■135

■231

■290

■128

Manjimup

■200

•175

•111

■101

•075

•049

•034

■0.54

■072

•124

-104

■151

■099

Afternoon readings for Cape Leeuwin taken at 4-0 p.m.

Tke Vegetation of Western Australia.

Ixxv.

TABLE V.— CLIMATIC ANALYSIS.

The first column of figures represents the average variation from the mean annual rainfall,
expressed as a percentage departure from the mean.

Humidity (four columns) —

The morning and afternoon readings for the four consecutive wettest, and the four con-
secutive driest months respectively (i.e., months of maximum and minimum pre-
cipitation).

Saturation deficit —

The first column gives the mean of the monthly readings for the year.

The second column gives the mean of the monthly mean readings for the four consecutive
wettest months.

The third column gives the mean monthly readings for the four consecutive driest months
{see Table II., in which these months are indicated).

MR = Meyer Ratio.

SP

— — = the mean precipitation for the four wettest months over the mean saturation deficit
SSD

for the same period.

Temperature —

MAT is the normal mean annual temperature.

MST is the normal mean temperatme for the four consecutive wettest months.

SP Seasonal precipitation r xu r xx x xl

3= i- L lor the tour consecutive wettest months.

MST Seasonal temperature

Staton.

Annual

Rainfall

Variation.

Humidity.

Sat

Def. (9

a.m.).

M.R,

(9-0

a.m.)

SP.

Temp. ° C.

SP.

Wet S.

Dry S.

Annual

Wet.

Dry.

SSD

MAT.

MST.

MST

Port George IV

17

68

49

102

26-1

30-0

37-2

Wyndham

20

58

50

42

38

•550

•444

•549

45

50

28-9

31-1

18-2

Derby

30

70

64

53

45

•492

•406

•534

51

53

27-6

30-3

18-0

Broome

32

70

64

53

45

•437

•364

•420

52

53

26-7

30-0

16-4

Hall’s Creek

29

48

32

35

28

•715

•658

-.559

27

24

25-6

29-4

13-7

Anna Plains

32

63

43

•601

•459

•491

24

24

Marble Bar

34

45

26

32

22

•690

•833

•768

19

11

27-8

33-1

6-9

Port Hedland

40

56

58

48

54

•518

•552

•606

23

15

24^4

30-6

6-8

Roeboume

43

49

40

39

33

•649

•734

•684

18

11

26-7

311

6-8

Mundiwindi

34

29

17

32

17

•634

•900

•462

16

5

22-8

29-4

5-4

Onslow

60

55

52

55

52

•486

•428

•579

19

13

24-4

24-4

5-9

Wiluna

35

40

29

41

30

•492

•659

•454

19

8

21^6

23-9

5-4

Meekatharra

35

40

24

33

20

•493

•688

•589

18

7

22-4

17-8

6-6

Cue

36

55

38

41

28

•435

•330

•520

20

12

9A • 9.

20-0

4-9

Laverton

31

51

37

47

34

•434

•326

•302

21

13

20-4

18-3

5-7

Menzies

30

55

38

39

25

•379

•280

•471

24

15

20-4

17-8

5'9

Rawlinna

29

63

36

39

23

•256

•200

•373

25

13

18-1

16-7

4-0

Carnarvon

37

63

59

56

60

•341

•225

•408

27

28

21-7

17^8

9-1

Yalgoo

28

72

50

47

30

•349

•122

•426

29

44

20-4

13-9

9-8

Kalgoorlie

22

64

43

47

30

•293

•211

•380

32

20

18^9

17-2

6-1

Southern C'ross

24

78

56

48

31

•267

•087

•460

39

61

18-0

10-0

12-5

Watheroo

74

39

•319

•113

• 537

51

95

17^7

14-2

24^0

Kellerberriii •

20

82

59

50

32

•228

•069

•408

58

113

18^0

12-2

16-2

Northam

20

83

61

59

36

•238

•072

•371

72

159

17-9

12-2

23-9

Esperance

14

76

64

62

63

•193

•114

•272

136

133

16'2

13-3

29-0

<Teraldton

19

68

59

55

60

•302

•179

•416

61

79

19-6

16-7

22^0

Katanning

17

84

66

57

37

•171

•060

•294

109

181

15-6

10-7

21-3

Perth Observatory

16

74

61

52

45

•246

•115

•383

141

214

17-9

13-9

45-2

Bunbury

16

78

69

59

57

•195

•095

•305

174

253

16-8

13-9

44-2

Dwellingup

87

66

62

40

•148

•046

•251

350

788

14-9

10^8

85-2

Cape Leeuwin

14

81

78

88

86

•085

•082

•087

411

313

15^1

14-4

45-3

Albany

14

79

69

71

66

•128

•077

•183

293

270

15 • 6

13-8

42-3

—

Ixxvi

C’harles Austin Gardner,

SUMMER RAINFALL (NOVEMBER-APRIL).

Isohyets in inches,

The Vegetation of Western Australia.

Ixxvii.

Plate II.

WINTER RAINFALL (MAV-OCTOBER).
Isohyets i;i iurlios.

Ixxviii.

Charles Austin Gardner.

I WHS, i&poO|i?’ri,6f.»»>,c I «,!/••

l^LATE III.

NORMAL MEAN TEMPERATCRE.

Isotherms in ° F for January.

The Vegetation of Western Australia.

Ixxix.

Plate IV.

NORMAL MEAN TEMPERATURE.

Isotherms in ° F. for July.

Ixxx.

Charles Austin Gardner.

Plate V.

RAINFALL VARIABILITY.

Variation in the annual rainfall expressed as a per cent, departure from the Mean.

The Vegetation of Western Australia.

Ixxxi.

Plate VI.

ilEYER RATIO.
/ P \

Ixxxii.

Charles Austin Gardner.

■ ?

Plate VTI.

The

Seasonal

Precipitation.

Seasonal saturation deficit

The Vegetation of Western Australia.

Ixxxiii.

Plate VITT.

The

Seasonal temperature
(Calculated in ° C. and mm.)

XXXl\’.

Charles Austin Gardner.

Plate IX.

THK THREK PKON’IXCES OF VEGETATIOX AXI) CLIMATE.

1. The Xoriheni Province . — Summer rainfall-winter drought. The four con.secutive
wettest months are Deoember-Maroh or January to April, the precipitation for
these four months being in excess of 175 mm.

'2. The South-West Province . — AVinter rainfall-summer drought. The four consecutive
wettest months are ^lay- August, the precipitation for this period being in excess of
1 75 mm.

JL The Eremean Province . — Rainfall for the four consecutive wettest months being less
than 175 mm. In the northern part (indicated by broken lines), the four wettest
7uonths are January- April. In the southern part (indicated by broken lines) the
four wette.st months are !\Iay- August. The middle region (unhatched) has its maxi-
mum pi-eilpitation between starch and June, the two wettest months being Marca
and June.

The \'egetation of Western Australia.

N9 J2I 123 t25 l27 129

'Ciuco.ir;'

Ixxxvi.

Charles Austin Gardner.

Plate X.

THE FORMATIONS OF THE VEGETATION.

^ . Tropical solerophyllous woodland.

' 2 . Monsoon woodland, savannah woodland, and riverain forest.

8. Savannah and open savannah woodland.

4. Triodia steppe,
o. Mulga Bush.

(1 Salt-bush steppe.

7. Scicrophyllous woodland.

8. Sand-heath.

9. Temperate savannah woodland.

10. Solerophyllous forest (“ Jarrah forest ”).

11. Mesophytic forest {Karri forest).

12. Desert.

The Vegetation of Western Australia.

Ixxxvii.

JOURNAL OF THE ROYAL SOCIETY
OF WESTERN AUSTRALIA.

Volume XXVIII.

1.— PERMIAN PEODUCTINAE AND
STROPHALOSIINAE
OF WESTERN AUSTRALIA.

By K. L. Prendergast.

Read 8th October, 1941; Published 20th April, 1943.
Communicated by Professor E. de C. Clarke.

CONTENTS.

Page

I. Introduction ... ... ... ... ... ... ... ... ... 1

II. Morphology and Bionomical Interpretation thereof ... ... ... 2

III. Trends in the Froductinae ... ... ... ... ... ... ... 8

IV. Classification ... ... ... ... ... ... ... ... ... 9

V. Description of Species ... ... ... ... ... ... ... 12

VI. Bibliography ... ... ... ... ... ... ... ... ... 57

VII. Plates ... ... ... ... ... ... ... ... ... ... 62

I. INTRODUCTION.

The correlation of beds in geographically distant localities has often
been based on the comparison of published lists of fossils. At many times
in the history of Australian Geology these comparisons have been made with-
out a re-examination of the specimens concerned :

“Creatures borrowed and again conveyed
From book to book — the shadows of a shade.”

The first specimens from Western Australia were described in 1883, and at
that time one specific name covered a multitude of forms which have now
been subdivided into several species. It is thus obvious that if the corre-
lations are to have any value, drastic revision of the naming of many
specimens is necessary. This paper, containing complete descriptions of the
Permian Productinae and Strophalosiinae of Western Australia is a small
contribution towards that revision.

The present time is opportune for this revision as the work of the
geologists of the Freney Kimberley Oil Co. and of Oil Search, Ltd., has
increased enormously our knowledge of the geology of the Permian and has
made available collections of fossils. I offer my thanks to Dr. Arthur Wade
of the Freney Kimberley Oil Co. and to Mr. H. Fletcher of the Australian
Museum, Sydney, for the loan of the specimens collected.

For the loan of specimens used in the preparation of this paper the
author thanks, as well as those already mentioned, Dr. H. M. Muir- Wood
of the British Museum, Professor E. de C. Clarke of the University of
Western Australia, Mr. A. G. Brighton of the Sedgwick Museum, Cambridge,
Mr. L. Glauert of the Western Australian Museum and Mr. F. G. Forman
of the Geological Survey of Western Australia. From Dr. Muir-Wood, Mr.
Brighton, Professor Clarke and Professor Wanner of Bonn I have received

K. L. Pre>?derGx\st.

much helpful advice. The text-figures have been drawn by Dr. D. H. Rayner
and the photographs prepared by Mr. A. Barlow of the Sedgwick Museum
and Mr. H, Smith of the University of Western Australia. The author
gratefully acknowledges her indebtedness to these persons and to many
others, too numerous to name, who have given her assistance.

The -work has been done while the author held a Hackett Studentship
of the University of Western Australia and an 1851 Overseas Scholarship.
She records here her thanks to the authorities concerned in these awards.

The publication of this paper has been made possible by a grant from
the Hackett Studentship Fund of the University of Western Australia.

The following abbreviations are used in the text : —

B.M., British Museum (Natural History).

Aus. Mus., Australian Museum.

W.A. Mus., Western Australian Museum.

U.W.A., Department of Geology, University of Western Australia.

G.S.W.A., Geological Survey of Western Australia.

The terminology as used throughout this paper is as defined by
Muir-Wood (1928). All measurements of specimens are given in millimetres
and the sign -f- after a number indicates measurement along a broken
specimen.

XL MORPHOLOGY AND BIONOMICAL INTERPRETATION

THEREOF.

External Features.

The general form of the shells of the Productinae and Strophalosiinae
is concavo-convex dorso-ventrally. There are, however, some exceptions.

Within the group Productus sensu lato .the concavity of the brachial
valve varies from strongly concave as in Krotovia to almost flat as in
Waagenoconcha. The pedicle valve is always convex and usually strongly so.
The curvature, as seen in longitudinal profile, may vary but little over the
whole profile or may change abruptly. In the latter case the shell is called
geniculate when the curvature has increased and fringed or flanged when it
has decreased. Fredericks has suggested a classification of the Productids-
based on the types of visceral cavity produced by the differences in curvature
of the two valves. He distinguishes three types.

A. Productus typici a. Pedicle valve with regular cimvature..

Brachial valve flat or gently concave. Visceral cavity deep..
e.g. Waagenoconcha imperfecta Prendergast.

B. Productus typici Pedicle valve with regular curvature.

Brachial valve concave, following the curve of the pedicle-
valve. Visceral cavity thin. e.g. Krotovia spimdosa
(Sowerby).

C. Productus prohoscidei. Pedicle valve regularly curved or

geniculate. Brachial valve geniculate. Development of
trail characteristic of the group, e.g. Productus productus
(Martin).

Permian Productinae and Strophalosiinae of W.A.

3

The terms suggested by Fredericks serve as useful names for the de-
scription of visceral cavities, but it is doubtful if they can be used as the
diagnostic feature by which to distinguish separate genera. The character
of the visceral cavity may or may not be a reflection of important anatomical
differences. It is, however, of little importance in modern lamellibranchs as
Pecten, for example, has one valve (usually the left) flat, while the right
valve may vary from slightly convex to rotund. There is here, within the
one genus, a great variation in the size of the visceral cavity. The example
chosen may not be strictly analogous, however, and where in a large number
of specimens the visceral cavity is of the same type, this, in combination
with other features, could certainly be used as a feature of diagnostic im-
portance.

In Aulosteges the shell has the same form as in Productus s.l. Where,
however, the area of the pedicle valve is high and reclined, this valve tends
to become flattened and the brachial valve is slightly convex, the form
approaching to the condition seen in the Orthotetince. Among the species of
Strophalosia the form of the shell does not vary beyond the limits already
described for Productus s.l. In Etheridgina the pedicle valve is often flat-
tened by attachment of the whole valve.

The area of attachment in Strophalosia and Aulosteges raises the ques-
tion of the position of the shell during life. In Productus s.l. the shell lay on
the larger pedicle valve with the brachial valve approximately parallel to
the substratum and the plane of symmetry vertical. The growth of a trail
allowed the animal to obtain its water supply from a level higher than that
■of the body. Possibly the shell sank by its weight into the substratum and
water taken into the body through the trail was cleaner and contained a
richer food supply. The trail would also serve as a protection to the animal
in providing only a narrow opening at some distance from the vital organs
and separated from them by a right-angled bend. In StrojAialosia the point
of attachment is at the apex of the umbo. It is not known for what period
of its life the animal was attached, but it seems unlikely, when the small size
■of the cicatrix is compared with the size of the shell, that this
condition of growth prevailed throughout life. However, whatever
the length of this period the shell must have gTown upwards from
the point of attachment at the umbo. In many Strophalosias, owing
to the geniculated or bevelled chai’acter of the interior of the brachial valve
the shell develops a small trail. Presumably, when the shell became too heavy
to maintain itself in the vertical position it fell on to its heavier pedicle
valve, the trail then serving as in Productus. An irregular deformation of
the umbo, presumably due to attachment, is sometimes seen in Aulosteges;
the same explanation may be offered in this genus.

The ornamentation of the Productince and Strophalosiince is essentially
spinose. Spines are developed most frequently on the pedicle valve and
rarely on the brachial valve which, where the pedicle valve is spinose, de-
vebps pits corresponding in position to the spines of the pedicle valve. The
spines may be of one or different sizes, they may be erect, oblique or adherent
while their position on the shell may vary from genus to genus. They may
be scattered irregularly or show a definite arrangement such as a row along
the cardinal margin or separating the ears from the body of the shell. They
may occur in concentric or radial rows or show a regular quincuncial arrange-
ment over the whole shell surface. The capacity to develop spines is probably

4

K. L. PRENDERGAST.

inherent in the animal, while the degree of their development will depend
upon external environmental conditions. Given the condition necessary for
spine development, that is, a plentiful supply of CaCO.i, any species will
probably develop spines in the same position in all its members. The inclina-
tion of the spines to the body surface will depend upon the hardness of the
sea-floor, arising at a high angle where the substratum is soft and being ad-
herent where it is hard. In modern lamellibranchs the temperature of the
water has a marked effect on the thickness of the shell, the Arctic forms
having a thick shell and the warm water forms of the same species a shell
thin almost to fragility. The difference between the Irwin forms, thin with
short spines of small bore and those from Mt. Marmion, thick-shelled with
heavy spines, of Tceniothcerus suhquadratus (Morris) is possibly due to the
difference in temperature of the sea at the two localities.

Both valves may be ornamented by ruga? and costff. The rugae are
formed at the growing edge of the shell and are thus in inverse positions in
the two valves. Costae are longitudinal folds perpendicular to the rugae.
Where both are present, the surface is reticulated as seen in the Dictyoclostus
group ; at the points of contact of the two series of folds an enlarged node
may be produced, sometimes giving rise to a spine. Wrinkles are more in-
definite than rugae, they may cross the visceral disc or be confined to the ears.
The lamellae seen in Strophalosia and Aidosteges are quite distinct from both
rugae and wrinkles. They are not folds but are the edges of the shell as it
was laid down in successive layers. The distinctness of these layers one from
another indicates that secretion of shell substance was not continuous but
intermittent and seasonal, a period of secretion being followed by a resting
phase. The control is a physical one but the response of the various genera
is dependent on the sensitivity of the animal itself and is, therefore, a bio-
logical factor.

In the structure of the cardinal margin these shells show a wide diversity
of form. In the Productus s.l. group a cardinal area is not ordinarily de-
veloped. In Dictyoclostus callytharrensis n.sp. however, and in some Indian
forms, e.g. Dictyoclostus indicus (Waagen) a narro-w concave “area” named
by Schuchert and Cooper a ginglymus is often seen on the pedicle valve. It
is to be noted that the ginglymus is only seen when the brachial valve has
been displaced. This ginglymus is a groove along the cardinal margin of the
pedicle valve into w’hich fits the narrow margin of the brachial valve hence
greatly increasing the efficiency of the articulation of the valves. In the genus
Produetorthis Schuchert and Cooper postulate that the ginglymus has arisen
from the degeneration of the area. This explanation cannot hold for Dictyo-
clostuSf as the early forms of the genus possess neither an area nor a gin-
glymus. Whether the ginglymus can, conversely, develop into an area, present
information does not allow us to state. It seems unlikely that development
would take this course unless teeth are present to take over the function of
the ginglymus as an aid to articulation.

Both Aulosteges and Strophalosia possess a true area though its form is
very different in the two genera. In Aulosteges it is linear in the earlier
species, but rapidly develops among the later species to a high plane triangle
produced ventrally and thus carrying the umbo of the pedicle valve away
from the brachial valve. In Strophalosia the area is much smaller, remains
linear throughout the genus and is produced over the hinge-line so that
the umbo of the pedicle valve approaches the brachial valve. A narrower
area is present in the brachial vah^e of Strophalosia.

Permian Productinae and Strophalosiinae of W.A.

.)

The area in both these genera possesses a distinct pseudo-deltidium and
in Strophalosia a pseudo-chilidinm in the brachial valve. The terms pseudo-
deltidium and pseudo-ehilidium have not the significance usually given them
in the description of embryonic brachiopods. The division of the area into
two lateral parts by the pseudo-deltidium is undoubtedly due to the inter-
ruption of the deposit of CaCOg along the cardinal margin by the projecting
ridges, the pseudo-deltidium being that part of the shell filling the space
between these ridges. At the places of formation of the ridges there is excess
CaCOg deposit and this projects slightly above and below the level of the
area, as Frederick's ‘^delthyrial ridges” above and where the downward
projecting ridges are more pronounced they take on the function of and are,
teeth. In Strophalosia the angle of divergence of the sides of the pseudo-
deltidium gives some indication of the angle of divergence of the roots of
the teeth though not necessarily of the teeth themselves as these may extend
laterally from their roots. That portion of the hinge-area, called, in the
brachial valve, the pseudo-ehilidium is the posterior root of the cardinal
process. The narrow grooves limiting its extent are the internal edges of
the sockets. In A.ulosteges, which has no brachial area, the posterior root of
the cardinal process is produced beyond the hinge-line as a flat triangular
plate, in the plane of the brachial valve, which partly fills the delthyriuin.
The pseudo-deltidium of Auhsteges may bear spines.

Internal Features.

Brachial Valve. — However the external appearance of the brachial valve
may vary on the inside it always has a flat visceral disc with possibly down-
turned margins. A medium septum arising at the base of the cardinal process
divides the shell into two halves. The septum varies in length from one-half
to two-thirds of the length of the visceral disc.

Muscle Impressions. — The adductor muscle impressions lie posterior to
the middle of the valve on either side of the median septum. They are den-
dritic in Aulosteges and Productus s.l. but non-dendritic in Strophalosia.
The presence of dendritic muscle impressions in Aulosteges and Productus
s.l. is a reflection of the poorly developed articular apparatus in these two
genera. Since the brachial valve is pivoted loosely on the cardinal process
it is capable of movement not only in a plane perpendicular to the surface
of the valve but also from side to side. This lateral skewing movement would
cause a differential movement within the muscle itself, the fibres nearer the
centre being less extended than the excentric fibres. A divided muscle would,
therefore, be mechanically superior. In examining a dendritic muscle it will
be noted that the bunches of fibres have an approximately radial arrange-
ment with the radii on the inside much shorter than those towards the lateral
margin. As the central muscles have a shorter pull this arrangment is what
would be expected.

In Strophalosia the muscle scars are non-dendritic but, except in
Wgndham.ia, are divided into posterior and anterior parts. The anterior
adductor is elongated in the longitudinal axis of the shell and lies very close
to the median septum while the posterior jiortion is elongated laterally, lies
above and close to the anterior adductor, but extends beyond it. The brachial
impressions arise at the lateral terminations of the posterior adductor im-
pressions.

c

K. L. Prendergast.

Brachial Impressions. — The brachial impressions are similar throughout
the group. In Strophalosia^ as already noted, they arise at the lateral
extremities of the posterior adductors and continue to the lateral margin at
an angle to the hinge-line varying from 4r)°-0°. They then run parallel or
neaz’ly so to the lateral and anterior margins to a point about one-third of
the width of the shell from the lateral margins when they curve through a
right angle towards the hinge-line. This line is followed until the impressions
are opposite the end of the median septum where they have another right
angle bend to meet at the anterior end of the median septum. In Productus
s.L and Aulosteges the brachial impressions arise at the anterior end of the
dendritic impressions and continue to the lateral border in a course parallel
to the hinge-line. When almost at the lateral margin they curve in a wide
arc and return to the base of the median se])tuni, again running parallel to
the hinge-line. In many specimens the returning arm is but faintly seen or
may be totally obscured. The interpretations of the brachial impressions
have been numerous. They are now described as the attachments of the
brachial arms which ])rojected ventrally from them. Muir-Wood (1928, p. 20)
has cited some evidence in favour of this view from the minute structure of
the impressions. The upward medial bend, where the impressions recede
from the anterior margin would correspond, then, to the pallial sinus of the
Lamellibranchs, being the region of water intake. The water would bo ejected
from the apices of the cones and this current would thus have no effect
on the basal attachments. The development of a fold in the brachial valve
with the corresponding sinus in the pedicle valve would serve further to
localise the ingoing and outgoing currents.

Cardinal Process. — In the description of the species the term ^Trifid’’
has been used to describe the cardinal process. To avoid misapprehension
that term is here defined as referring to a cardinal process which is bilobed
in ventral aspect and trilobed in dorsal aspect or which is a modification of
this type. The dorsal and ventral aspects are the views of the cardinal
])rocess as seen from the brachial and pedicle valves respectively.

To expand this definition the cardinal process of Aulosteges wangenheimi
(Vern.) may be considered (refer to Fig. 5, p. 34). In this species the
cardinal process is perpendicular to the plane of the brachial valve and is
supported by two lateral ridges arising from the cardinal ridge. These
ridges, which are separated medially by a deep sulcus, give rise to two
convex masses, the lobes of the ventral aspect, separated by a depression.
The dorsal aspect shows the structure of the process clearly. It has the
form of a double-S. The median convexity of this double-S is the central
lobe, and the end arms the lateral lobes of the dorsal aspect while the two
concavities are the lobes of the ventral aspect. The whole process has the
form of an open and indented cone with its apex at the hinge-line. The
growth-lines of the structure may be followed round the dorsal face and
are ])arallel to the base of the cone.

AH the cardinal processes examined were modifications of this type.
Thus in Aulosteges haracoodensis Eth. fil. the process is not in such an
advanced stage of development. The dorsal aspect shows the process elon-
gated with the lobes becoming distinct some distance from the hinge-line.
The ridges of the ventral aspect have remained as ridges and are not enlarged
into the convexities seen in Aulosteges wangenheimi (Vern.). They are
indented posteriorly having a quadrilobed termination.

Permian Productinae and Strophalosiinae of W.A.

7

The variations within Productus s.l. are numerous. In the semiretieulate
group as Muir-Wood (1928, p. 18) says; —

Cardinal Process of Aulosteges barncoodensis Eth. fil. (X3).

*‘The cardinal process projects beyond the margin of the brachial
valve and is divided by a median furrow into two parts which are smooth
and rounded ; these are continued dorsally as to two curved laminae,
separated from the median septum by deep sulci. The dorsal view of
the process show's it to be trilobate and transversely striated.”

This condition can be readily understood by reference to Fionre 2 (p. 15)
showing the cardinal process of Dictyoclostus callythorrensis n.sp. In this
species as in all Dictyoclostids the whole cardinal process is flattened and
widened and the lateral lobes have been rotated through nearly 90° until,
instead of being parallel to the central lobe, they lie at right angles to it
and parallel to the hinge-line.

In Taeniothaerus, Linoproductus and Waagenoconcha the modification
has taken quite a different form, as in these genera the tendency has been
for the process to elongate and for the lateral lobes to be distinct and
separated from the central lobe (see Taeniothaerus subquadratus (Morris),
Figure 3, p. 28). The median depression of the ventral aspect is distinct
and continues to the apex of the process but the posterior part of the process
is now trifid even in its ventral aspect. In Taeniothaerus the process is still
compact, but in Linoproductus (e.g. Linoproductus cora (d’Orb.) Kozlowski,
P- 14) the lateral lobes have splayed out and the process appears;
winged. In Waagenoconcha (e.g. Waagenoconcha humholdti (Waagen),
Kozlowski, fig. 9, p. 15) the dissection of the lobes is continued further
towards the hinge-line than in the other g'enera. In the aged specimens
Kozlowski found that the median lobe sometimes divided giving what has
been described as a quadrilobed cardinal process.

The shells so far considered have had the median septum separated
from the cardinal process by a sulcus of varying depth. In the Strophalosias,
on the other hand, the median septum arises Avithin the cardinal process. The
septum swells out betAveen the sockets and completely fills the median
depression betAveen the ventral lobes. This sAvelling is showm in Strophalosia
Umberleyensis n.sp. (Fig. 8, p. 48) in Avhich a part of the depression may
be seen posterior to it. This outgrowth of the median septum would give
additional support to the socket plates. The median septum in Strophalosia
etheridgei n.sp. (Fig. 7, p. 44) is not unduly swollen, but has, nevertheless,

8

K. L. Prendergast.

completely obliterated the median depression. On its dorsal face, also, the
process of this species is much reduced, faint depressions marking the posi-
tions of the dorsal concavities.

Where the median depression is present in the ventral aspect, as in
Anlosteges and Prodnetus s.l. the depth of this depression is regulated to
some extent by the inclination of the cardinal process to the plane of the
brachial valve. Where the process is erect the depression is shallow but
where it becomes sharply bent over, as in Anlosteges ivangenheimi, the sulcus
is very deep.

Marginal Eidgesy etc. — Marginal ridges are sometimes developed in
species of this group, for example, Strophalosia kimberlegensis. These ridges
are always low and not prominent. In some species of Aidosteges a pro-
nounced ridge may be developed along the hinge-line, as in Anlosteges
wangenheimi (see Fig. 4, p. 33). This ridge is placed anterior to the hinge-
line.

Pedicle valve . — The muscle impressions of the pedicle valve are of the
same general form although they vary throughout the group. The adductors,
dendritic in Atilosteges and Prodnetus s.l., non-dendritic in Strophalosia, are
median and separated by a groove or a small ridge. The diductors are large,
flabellate and longitudinally striate. In some species of Strophalosia the
ears of the pedicle valve are thickened so that when the shell is closed the
ears fit closely one upon the other. This thickening is seen as a flat ledge
on the inside of the valve and projects slightly over the visceral cavity.

For a description of the microscopic structure of the shell reference
should be made to Muir-Wood (1928, p. 29) and Dunbar and Condra
(1932, p. 179).

TIL TRENDS IN THE PRODFCTINAE.

Although sufficient data are not yet available for the recognition cf
lineages, nevertheless some indication of possible trends can be given, based
on the fauna of other areas, and these ideas can be applied to the Australian
forms.

The table below shows the distribution and characters of the species of
Anlosteges from the Permian of the Glass Mountains, Texas (King, 1930).

—

Wolfcamp.

Hess.

Leonard.

Word,
Delaw’are
Mtn, White
Limestone of
Guadalupe.

A. wolfcampensis
A. rnedlicottianus

lb, 2b, 3b
lb, 2b, 3b

lb, 2b, 3b

lb, 2b, 3b

lb, 2b, 3b

A. ynagnicostatus
A. triagonalis
A. subcostatus
A. beedei
A. guadalupensis
A. tuherculatus

lb, 2b, 3a
la, 2b, 3a

lb, 2b, 3a
la, 2b, 3a
la, 2a, 3b
la, 2a, 3a

lb, 2b, 3a
la, 2b, 3a

la, 2a, 3a
la, 2a, 3a
la, 2a, 3a

Where 1. radial costae ; b, present, a, absent.

2. concentric wrinkles ; b, present, a, absent.

3. area ; b, low and linear, a, higher and triangular.

Permian Productinae and Strophalosiinae of W.A.

9

As may be seen from the table the shells change from a lirate, concen-
trically wrinkled form with a low linear area to an unwrinkled form with
a high triangular area ornamented only by spines and growth lines.

In India, the evidence of a parallel evolution of Aulosteges is not sa
clear, but is indicated. Aulosteges medlicottianus occurs in the lowest fossili-
ferous beds, the Amb, of the Salt Range, while Aulosteges (Strophalosiina)i
tibeticuSj which has the same characters as Aulosteges magnicostatus (con-
centric wrinkles with radial costae anteriorly) is found in the Middle Pro-
ductus limestone. Aulosteges dalhousii from the Upper Productus Limestone
has the same characteristics as Aulosteges guadalupensis.

Among the Australian form a similar parallel can be traced. A series
of forms appears connecting Aulosteges haracoodensis with T aeniothaerus
sub quadrat us. The latter species has no concentric wrinkles, a low linear
area and the spine bases on the anterior part of the shell sometimes elon-
gated as radial ridges. Aulosteges haracoodensis is ornamented by spines
and growth lines only, and has a higher triangular area. Both these end
forms occur in the Fossil Clift Beds, Irwin River, although in the North-West
Basin Aidosteges haracoodensis does not appear until much higher in the
sequence. Judged by the rate of change of the American forms T aeniothaerus
suhquadratus would appear in the equivalent of the Hess beds and Aulosteges
haracoodensis not until the Word. The evolution of the forms may, how-
ever, have taken place at different rates in the two areas.

A second shred of evidence of the general stage of development of the
Western Australian fauna is provided by the Dictyoclostus group. In that
genus there is a marked tendency for the area of reticulate ornament to
decrease as the species become younger. Thus in Dictyoclostus semireticidatus
from the Lower Carboniferous of the British Isles the reticulation may be
found over the whole shell though not as clearly marked anteriorly. In the
Permian species, however, the reticulation is confined to the posterior part
of the shell. D. spiralis, indicus, vishnu, aratus and stihcostatus is an ascend-
ing stratigraphical series from the Salt Range (Waagen 1884) and shows
a gradual decrease in the area of reticulation of the species in the order
named. The Western Australian shells, Dictyoclostus callytharrensis, have
the reticulation covering but a small area of the valve and possess a ginglymus
while the ears are separate from the flanks. All these features are seen only
among the younger members of the genus.

IV. CLASSIFICATION.

Family PRODUCTIDAE Gray, 1840.

In his original description of the family. Gray included the genera
Productus, Strophalosia, Chonetes, Leptaena, Orthis, Strophomena and
Calceola. King (1846, p. 94) restricted the family to those shells “distin-
guished by the form of the ovarian spaces and the presence of spines, and
excluded the genera Strophomena, Orthis, Leptaena, Chonetes and Calceola,
Gray (1848, p. 438) was opposed to this restriction and enlarged the family
to its original content. Gray^s opinion, however, was not widely accepted,
and King's diagnosis became the diagnosis of the family.

10

K. L. PRENDERGAST.

It was accepted thus by Waagen (1884, p. 611) who said: —

“the most striking- character is the existence of reniform prominent
ridges on the inside of the dorsal valve, a character which in this
strong and distinct development, is entirely restricted to the
Productidae.^'

Waagen subdivided the forms so characterised into two sub-families,
Chonetinae and Productinae.

The Chonetinae were characterised by the presence of teeth and a car-
dinal area in the pedicle valve and of non-dendi’itic muscle impressions. This
sub-family included the genera Chonetes^ Strophalosia^ Chonetella and
Daviesiella, Most of these forms are free.

The Productinae were defined as Productids in which the pedicle valve
is without cardinal teeth and the adductor muscle impressions are dendritic.
Some of the forms are fastened by the entire pedicle valve or by spines.
Waagen placed the genera Aulosteges, Productella, Productus and Margini-
fera in this sub-family.

Schuchert (1913, p. 389) changed the emphasis in King's diagnosis
and defined the Productidae as “Strophomenacea with hollow anchoring
spines."

Schuchert also altered the content of the sub-families by using the means
of attachment of the shell as the main diagnostic feature. Thus the
Chonetinae became

“Productids with a few anchoring spines restricted to the ventral
cardinal margin,"
and the Productinae

“Productids with the anchoring spines more or less abundant over
the entire ventral and sometimes also over the dorsal valve,"
while he founded a new sub-family the Strophalosiinae to include

“Productids anchored to foreign objects by spines or by most of the
ventral shell.”

Such a division of the Productidae is purely arbitrary and while it groups
together forms superficially alike, makes no allowance for a similarity of
external form due to growth under similar environmental conditions. The
sub-familv Strophalosiinae^ for example, includes the genera Chonopecttis^
Strophalosia, Aulosteges and Etheridgina.

Strophalosia King, the type genus of the sub-family is an attached
form, being cemented by the umbo of the pedicle valve and/or by spines. It
has a welf-developed area on each valve, non-dendritic muscle impressions
.-and strong teeth. Etheridgina possesses an area only on the pedicle valve,
teeth may be developed and the shell is cemented directly by part or the
whole of^the pedicle valve and by spines. The muscle impressions are as in
Strophalosia.

The genus Chonopectus Hall and Clarke is represented by a single
species Chonopectus fisheri Norwood and Pratten from the English River
beds of the Kinderhookian (equivalent to K beds. Lower Carboniferous) of
Burlington, Iowa, U.S.A. This species is essentially a small Chonetes with
a distinctive ornamentation and attached at some period during its life
history. It is a restricted form, both geologically and geographically.

Permian Productinae and Strophalosiinae of W.A.

11

Aulosteges Hclmersen is distinct from Strophalosia in the possession of
dendritic muscle impressions. The pedicle valve carries an area, usually
high, while that of the brachial valve is small or absent. The articulation is
weak, the pedicle valve being without teeth.

It is obvious, then, that within his sub-family Strophalosiinae Schuchert
has included forms very different in structure. Its attached habit seems an
insufficient reason for the separation of Chonopectus from those forms, the
Chonetinaej to which it is closely related, and Schmidt's placement of it
(1929, p. 21) in that sub-family must be regarded as the more correct.
Aulosteges^ both in the possession of dendritic muscle impressions and in
the pattern of the brachial impressions, approaches more closely to the
Productinae than to Strophalosia and its replacement in the Strophalosiinae
would seem a more natural position.

It is admitted, however, that little is known of the derivation of
Aulosteges and Strophalosia, Licharew (1934 (a), p. 509) has, on the
ground of insufficient evidence for their separation, abolished the sub-families
and included all the genera in Prodiictidae. He reverts to King's diagnosis
and defines the family as

^‘Shells free, attached by spines or cemented directly. Dorsal valve
flat or concave. Cardinal margin straight. The whole surface or only
the shoulders of the ventral valve bedecked with hollow spines. Cardinal
process prominent. Dorsal valve with reniform brachial impressions."

To Productus sensu lato, Chonetes sensu lato, Strophalosia^ Productellay
Aulosteges and Etheridgina he adds Teguliferina and Scacchinella. Dunbar
and Condra (1932, pp. 189, 191) and Diener (1927, pp. 30 and 31) also in-
cluded these genera in the Producitidae.

Scacchinella Gemmellaro and Teguliferina Schellwien are the genera
intermediate in character betiveen Aulosteges and Productus respectively and
the Richthofeniidae. These genera, however, possess characters which would
ally them to the Richthofeniidae rather than to the Productidae. Scacchinella
and Aulosteges possess in common the high produced area (higher and more
produced in Scacchinella) and the method of attachment by the tip of the
area. Scacchinella is quite distinct, though, in the possession of a well-
developed median septum in the pedicle valve and in the character of the
cardinal process with its supporting rods. In these characters it approaches
Richthofenia, Teguliferina in which the productoid characters are still
recognisable, resembles closely the Richthof eyiiidae in its mode of growth and
in the operculiform character of the brachial valve. It seems reasonable tO'
assume that the peculiar characters of Richthofenia are in part a result of
its mode of growth and therefore that Scacchinella and Teguliferina^ as the

possible forerunners of a group so distinctive as the Richthofeniidae, should
be included in that family.

A different method of approach to this subject of the classification of
the Productidae has been used by Sutton (1938), with astounding results.
As he points out, the shape of the visceral cavity as a diagnostic feature
has been given prominence in Productid literature although no author has
attempted a classification based upon that feature. Sutton has. He divides
the Mississippian Productidae into two sub-families, the ProductelUnae and
the Productinae. The ProductelUnae have a thin visceral cavity of the type
described by Fredericks as Productus typici and include the genera

12

K. L. Prendergast.

Leptalosia, Strophalosia^ Productella, Gigantella and Productma Sutton.
The remaining Productid genera of the Mississippian rocks are grouped as
the Productinae and possess in common a ‘darge and medium to deep visceral
cavity — Frederick’s Productus typici cc P

This cannot be regarded as a biological or palaeontological classification;
it is as Sutton (p. 538) says, ^‘a classification by which different forms may
be recognised,” in other words, a key, and a key only for the Mississippian
Productidae. Sutton supports this ‘^classification” by reference to one now
being worked out by Stoyanow, who is using as a diagnostic feature the
character of the cardinal process, whether trifid or bifid. Examination of
the Australian Productids shows that all have essentially the same type of
cardinal process, but Stoyanow will doubtless define his terms more accur-
ately when his paper is published.

In the classitication which follows the author has tried to group closely
related forms together, and finds that in so doing he has reverted largely to
AYaagen’s original diagnoses of the sub-families while following Licharew in
the definition of the Productidae.

Family Productidae Gray 1840. Synopsis of the contents of the British
l\rusemn. 42nd Edition, p. 151.

Shells free, a+tached hy spines or cemented directly by pedicle valve.
Cardinal margin straight. Surface ornamentation of spines over whole or part
of shell. Brachial valve with reniform brachial impressions and median septum.
Cardinal process prominent.

Sub-family 1. CltonefiTiae Waagen 1884. Salt Range Fossils, I. Productus-
Limostone Fossils: TV (fasc. 3) Brachiopoda, Pal. Ind. Ser. XIII, p. 612.

Productids with large spines only along cardinal margin of pedicle valve.
Cardinal area and teeth in })edicle valve. Muscle impressions non-dendritic.

Genera. Chonctea sensu lato.

Chonopectus.

Daviesiella.

Sub-family 2. Productinae Waagen 1884, op. cit., pp. 612, 613. Productids
with dentritic muscle impressions and without teeth. Hollow spines present
over whole or part of shell.

Genera. Productus sensu lato.

Productella.

Aidosteges.

Sub-family 3. Slrophalodinae Schuchert 1913, in Zittel K., Textbook of
Palaeontology edited by C. R. Eastman, 2nd edition p. 391.

Productids attached by spines and by part or whole of the pedicle valve.
Ornamentation usually spinose. Muscle impressions non-dentritic. Pedicle valve
with cardinal area and teeth.

( Jenus. Strophalosia.

Heteralosia.

V. DESCKIPTION OF SPECIES.

DICTYOCLOSTUS Muir-Wood emend.

1930. Muir-Wood, A7in. Mag. Nat. Hist. (10) Y p. 103.

GeU'C-type. — Original designation. Ano^nites semireticulatus Martin (in part).
Petrifacta Derl)iensia, Wigan 1909, p. 7, pi. xxxii, figs. 1, 2; pi. xxxiii, fig. 4.

Diagnosis . — Shell elongate or quadrate in outline, hinge wide; ginglymus
may be present. Pedicle valve convex or geniculate ; visceral disc with semi-
reticulate ornamentation; trail costatc, costae may bear spines both on trail

Permian Productinae and Strophalosiinae op W.A.

13

and visceral disc; rows of spines on ears and cardinal slopes; diaphragm
absent. Brachial valve concave or geniculate; marginal ridges prominent,
extending along hinge. Hinge teeth and sockets not developed.

Range, — Carboniferous and Permian.

Remarks. — The diagnosis has been emended to include semireticulate
productids with a ginglymus (see p. 9). The Western Australian, Timor and
Indian species Avhich would normally be placed in this genus are sometimes
found to have a ginglymus though agreeing in other particulars with species
of this genus. Where the margin is interrupted by the cardinal process, the
ginglymus is notched to give the “delthyrium,” into which the cardinal process
fits, as previously suggested by Dunbar and Condra (p. 18). Below this
notch and on each side, thickened ridges which gradually die out laterally,
articulate with the transverse ridges of the brachial valve. As would be
expected, the ginglymus is not seen if the two valves fit tightly, but is on1v
exposed when the brachial valve is slightly displaced. This was proved in
a specimen (S.M. No. 3604) of Dictyoclostus indicus (Waagen) from the
Salt Range; it possessed both valves, and was without an area, but showed
an internal groove in that position when the brachial valve was removed.

The ginglymus has been noted in one other genus. Productortlns ot‘
Schuchert and Cooper (1932, p. 83). In Vroductorniis, Schuchert and Cooper
believe that the ginglymus has arisen from the reduction of the area. In
Productids it seems more likely that the area arose from the ginglymus.

The Permian species of Dictyoclostus, as far as can be seen from a study
of the literature, differ from the Lower Carboniferous species in the restric-
tion of the reticulation to the posterior part of the shell, the separation of
the ears from the flanks and the possession of an articulating ^^area.’’

The larger Western Australian members of the genus Dictyoclostus are
undoubtedly closely related, probably in the same way as the spiralis-
subcostatus grou]> of the Salt Range, and it is only lack of sufficient material
which prevents the apjn’eciation of their relationships.

Dictyoclostus callytl: arrensls sp. nov.

PI. i, figs. 1-7, PI. ii, fig. 1.

1903. — Troductns semireticulatus Martin, Etheridge, jun., p. 18, pi. ii.
figs. 3-0.

1907. — Produci-us semireticulatiis Martin, Etheridge, jun., p. 29.

1910. — Producius semireticulatus Martin, Glauert, p. 87.

1924. — Prodnetus .^emireticulaius Martin, Chapman, p. 36, sx)ecimens Nos.
2746, 2748.

1931.- — Productv.‘^ semireticulaf-us Martin, Hosking, pjn 8, 22.

Types.

Ryntypes. — G.S.W.A. 1/4967 (b), i/j mile west of Callytharra Spring, Woora-
mel River District ; Callytharra Limestone.

Paratypes. — G.S.W.A. 1/4967 (a), same locality and horizon.

Other specimens. — IT. W.A. 12400, Fossil Cliff, Irwin River; Fossil Cliff
horizon.

G.S.W.A. 1/4668, south bank of Wooraniel River, mile above Callytharra
Spring; Callytharra Limestone.

G.S.W.A. 1/4654.

Aus. Mus. F 36507, 36508, Barragooda Pool, Arthur River, North-West
Division. Callytharra Stage.

Aus. Mus. F 36237, Jimba Jimha Station, left bank of Gascoyne River,
near Winnemia, North-West Division. Byro Stage.

Aus. Mus. F 38147, 1% miles west of Williambury-Middalya Gate, Middalya
Station. Callytharra Stage.

14

K. L. Prendergast.

Diagnosis . — Shell large^ semicircular in outline, greatest width at hinge-
line; ears large and reflexed, offset from flanks, flanks convex to steep.
Pedicle valve sinuate, regularly curved through 270°. Visceral disc occupying
about half curvilinear length of valve, rugae and costae regular and equally
prominent on visceral disc, costae becoming irregular on trail, two or more
coalescing to form irregular folds; rugae absent on trail. Small spines,
quincuncially arranged, arising at intersection of costae and rugae, with
larger spines, more widely spaced, on rest of shell. Row of spines between
ears and flank and row along cardinal margin.

Brachial valve geniculate; ornamentation as in pedicle valve.

The size of the specimens is shown by the table : —

Syntypes.

Paratypes.

Largest spec.,
Irwin R. —
12400.

Length of hinge-line

59

59

58 44 +

49 +

Height

37

38

33+ 33 +

46 +

Curvilinear length

70

71

50 50

95

Shells too crushed to allow measurement of thickness.

Description of the syntgpes . — Shell semicircular in outline with the
hinge the greatest width of the shell.

The pedicle valve is curved through an angle of 270°. The transverse
curve of the valve is a high arch with almost vertical sides and a shallow
median depression. A shallow sinus arises at about the middle of the
visceral disc and continues forward^ becoming shallower and Anally disappear-
ing on the trail thus leaving the margin of the valve entire. The ears are
large and reflexed; each is separated from the body of the shell by a sulcus,
and this is flanked by a fold bearing a roAV of spines (3-4). A second row
of spines runs parallel to the hinge-line and close to it. The semireticulation
is marked on the visceral disc, covering 35 mm. along the curvilinear length,
the costae and rugae being equally prominent. The costae cannot be traced
to the tip of the umbo, where they have been probably I'emoved by weather-
ing, but traces of them can be seen on the ears. They increase in number on
the visceral disc botJi by division and by intercalation ; no increase takes
place on the trail and the costae lose their height, remaining scarcely elevated.
The rugae are closely spaced near the umbo, becoming farther apart
posteriorly. The spines on the visceral disc are small and oblique and
arranged in approximate quincunx, each spine arising from a node at the
intersection of a ruga and a costa. Spines do not arise from all the nodes.
As the spines are regularly spaced and the number of costae increases an-
teriorly, there is some irregularity, but every fourth to sixth ruga bears
spines, these being separated by eight or more costae. The larger spines on
the trail also tend to quincuncial ari’angement. Below each on the trail is a
fold which replaces the two or more costae above each spine, this folding
being independent of the costae, though two or more are absorbed on each
fold.

Permian Productinae and Strophalosiinae of W.A.

15

The brachial valve is strongly geniculate. The ornamentation differs
from that of the pedicle valve only in the absence of spines. The pits, which
replace the spines on this valve, are not placed opposite them, but appear
anteriorly. This is particularly noticeable in the row of pits on the sulcus
separating the ears. No trace of a cardinal area has been observed on this
valve.

The interior of the valve is not seen on the syntypes.

Description of the paratypes — Internal features, — The paratypes show
the gingiymus more clearly than either of the syntypes. It is divided by a
wide triangular notch. The cardinal process of the brachial valve projects
but a little above the level of the hinge as a truncated triangular ridge.

Internally the muscles of the pedicle valve are those characteristic of the
genus, longitudinally striated diductors enclosing dendritic adductors, the
place of insertion of the muscle much thickened.

Variation within the species. — The topotypes of this species show marked
variation in the depth of the sulcus between the visceral disc and the ears.
When the sulcus is shallow and not pronounced the rugae extend on to the
ears so that these are semireticulate. This ex,tension of the semireticulation
has an effect on the size of the auricular spines, since where the rugae are
present the spines are small and similar to those of the visceral disc, these
replacing the larger spines seen in specimens with a deep sulcus. Recognising
this as a variation within the species it is then possible to include the two
forms distinguished by Hosking (1933, p. 46) in the one species.

A second variable feature of the specimens is the extent of the semi-
retieulation along the curvilinear length of the shell. Although owing to the
crushing of many of the specimens it is difficult to determine the original
shapes, it would seem that rugae are not developed after there is a change in
the direction of growth, even where this does not amount to a geniculation.
This relation between the shape of the shell and the ornamentation probably

Figure 2.

Cardinal process of D. callytharrensis sp. nov. (x6).

16

K. L. Prendergast,

exists throughout the semiretieulate group. Dictyoclostus semireticulatus
(Martin), for example, has semiretieulate ornamentation throughput its
growth, and no change takes place in the curvature.

The Irwin River specimens show the same range of variation as those
from the type locality.

Comparison with other species. — The specimens of this species have
little in common with Productus (Dictyoclostus) semireticulatus Martin s.
str. to which species they have been previously assigned. They ditfer in
having the brachial valve geniculate, the rugae restricted to the visceral disc,
the arrangement of the spines on the visceral disc and the division of the
ears from the flanks. Briefly the only characteristic common to both species
is the presence of semiretieulate ornamentation.

The species has much in common with Upper Carboniferous and Permian
forms of India and China. It probably occurs in Timor (Basleo) as Broili’s
figures and specimens (1916 pi. cxvi figs. 14-16) show the same characteris-
tics as the Australian species and Broili gives a reference to Etheridge (1903)
in his synonymy. Of the Chinese specimens P. taiyuanfuensis Grabau as
figured by Chao (1927 p. 30 pi. i fig. 10; pi. ii figs. 1-12; pi. viii fig. 16) from
the Taiyuan Series is very close to 1). callytkarreivsis. The costation of the
Chinese shells is much coarser (the costae vary from 0.5 to 1.3 mm. in width
as compared with 0.3 to 0.5 mm.), the larger spines are more numerous and
more closely spaced, particularly the I’ow along the inner edge of the ears
where each spine touches its neighbour.

Z>. callytharrensis is undoubtedly related to the P. sj^iralis-subcostatus
group from the Salt Range. Freeh first suggested that the forms P. spiralis
and P. indicus were possibly the younger and older specimens of the same
species, and Broili supported this. More lately Cowper Reed (1931 p. 2) has
suggested that the whole group, P. spiralis, indicus, vishnu, aratus and
subcostatus are probably all members of one species group. The neax’est form
of this group to P. callytharrensis is P. indicus, but the Western Australian
specimens may be distinguished from that species by the tendency in the
Indian species for the costae to converge towards, and disappear in, the
sinus. Also the spiral arrangement of the costae on the ears cannot be seen
in the Australian specimens. The costation is much coarser in P. indicus
(7 costae in 10 mm., 10 mm. from the umbo).

Dictyoclostus callytharrensis var. wadei va.r. nov.

PI. II., figs. 2-4.

Bolotype. — U.W.A. 20453, fernigiiious limestone, two miles east 10° south
of Mount Nicholson, Kimberley Division; Nooneanbah Series.

Topotypes. — U.W.A. , A.58, A. 59, A.65, same locality and horizon.

Diagnosis. — Shell as in Dictyoclostus callytharrensis but -with approxi-
mately hexagonal outline and arched umbonal region. Ears reflexed but not
offset from flanks. Pedicle valve sinuate, with semireticulation on almost
one-third of curvilinear length of valve, costae irregular, prominent and
enlarged on trail, converging towards sinus. Brachial valve unknown.

Description. — The holotype of this variety is a weathered and decor-
ticated specimen. It is 63.7 mm. wide at the hinge-line and 54.5 mm. long.
The semiretieulate ornament of the visceral disc is badly worn, but probably
occupied about 34 mm. along the curvilinear length. The ears, though refiexed,
are continuous with the anterior margin of the valve. The trail is ornamented

Permian Productinae and Strophalosiinae of W.A.

IT

by large costae, variable in width, bearing large spine bases; these show
no regular arrangement. The costae tend to approach and disappear in the
sinus. Two large spines are on the ears and three on the flanks just above
the angle separating the ears. There is no enlarged rib or hollow on which
they are placed. The umbo is rounded and barely overhangs the hinge-line.
The ginglymus, if present, is obscured by matrix.

Variation within the species. — Owing to the paucity of specimens (5)
it is impossible to determine the limits of variation.

Comparison with other species. — This variety differs from D. callythar-
rensis in the shape of the ears (here continuous with the anterior margin
but in D. callytharrensis distinct and joining the lateral margin about halfway
down the flanks), the greater convexity of the flanks and the invariable
appearance of a well-marked sinus in the pedicle valve. The folds, too, in
this species are enlarged costae and not formed, as in D. callytharrensiSj by
two or more costae coalescing. In all these particulars they approach P.
indictis AVaagen, but continually differ in the much finer ornamentation.

It is possible that this form should constitute a distinct species, but in
the absence of any definite knowledge of their stratigraphical relations to
D. callytharrensis I have preferred to designate them as a variety. They are
obviously closely related to I), callytharrensis mihi.

Dictyoclostus gratiosus (AVaagen).

PI. ii, figs. 5-7.

1884. — Producius gratiosus AVaagen, p. 691, pi. Ixxii, figs. 3*7.

1893. — Prodiictus gratiosus AVaagen, Rothpletz, p. 76, pi, x, fig. 15.

Idld.—Productus gratiosus AA^aagen, Broili, p. 12, pi. cxvi, figs. 4, 5, 7-13.

1927. — Prodiictus gratiosus AA^aagen, Chao, p. 44, pi. iv, figs. 6-10.

1928. — Producius gratiosus AVaagen, Hamlet, pp. 19, 20.

21aterial. — Aus. Mus.

F 37567 — Vi mile east of shale outcrop, Minilya River, North-AVest Division,
Wandagee Stage.

F 37568 — 14 mile west of shale outcrop, same locality and horizon.

F 37569-37571 — Bank of Minilya River, same locality and horizon.

Diagnosis . — Shell small to moderate in size, outline hexagonal. Pedicle
valve inflated, geniculate; brachial valve concave. Greatest width at hinge-
line, ears small and pointed.

Pedicle valve with flat visceral disc, shell geniculate. Ornamentation
reticulate on visceral disc, costate on trail. Strong sinus. Small erect spines
scattered on trail and alar extremities. Costae converging towards sinus.

Brachial valve concave with median fold. Ears excavated, smooth or
lightly costate. Convergence of costae toAvards fold. Valve with reticulate
ornamentation on posterior part of shell.

Internal characters not knoAvn.

Description.

Dimensions.

Length of hinge-line

27-8

18-0-h

16-6 +

25-0

14‘5-h

Height

Curvilinear length of pedicle

17-7

13-7

16*1

15-8

11*7 +

valve

30*9

21-0

32*3

31*8

21-0

18

K. L. Prendergast.

The shells are small to moderate in size as may be seen from the table
of dimensions. They have an irregularly hexagonal outline when viewed
from the pedicle side, with the hinge-line the greatest width of the shell.

The pedicle valve has a fiattish visceral disc which, following the geni-
culation of the shell, passes into a regularly curved trail. Transversely the
shell arch is high, indented medially by a strong median sinus and falling
gradually to the lateral margins in a slightly convex slope. The umbo is
small, pointed and slightly overhanging the hinge-line. The ears are small
and pointed, they have a convex profile in longitudinal section. The orna-
mentation is reticulate on the visceral disc, the reticulation sometimes irregular
due to the enlarged rugae on some specimens. The rugae do not develop
anterior to the geniculation, the trail carrying costae and scattered spine
bases. The latter are rarely numerous, not more than 4 or 5 being present.
The co.stae converge slightly towards the sinus giving a very distinct and
characteristic appearance to the shell.

The brachial valve is trapezoidal in outline, the long straight hinge-line
and the anterior margin being the parallel sides. It is regularly concave
with a median fold which expands towards the anterior margin. The ears
are excavated to fit tightly with those of the pedicle valve. The ornamenta-
tion is reticulate over the jDOSterior third of the shell, the rugae then develop
irregularly and the ornamentation of the rest of the shell may be described
as costate. The costae radiate from the umbo, those in the central part of
the shell converging towards and finally coalescing on the median fold.

The internal features of this s]iecies are not shown by any of the
Western Australian specimens.

Comparison with other species . — This species is distinct from other
Western Australian scmireticulate productids in its small size. The converg-
ence of the costae towards the sinus in the pedicle valve and the fold in the
brachial valve serve to distinguish it from other small members of the semi-
reticulate group.

In the Western Australian specimens the convergence of the costae is
not so marked as in some of the Indian and Timor forms. Examination of
the Timor specimens shows, however, that there is a large range of variation
in the extent of the convergence.

Remarks . — As the internal features of this species are not known the
generic placement is insecure. In external appearance the shells are similar
to Marginifera Waagen, and knowledge of the internal structures is neces-
sary before it can be said that they do not belong to that genus.

Dictyoclostus spiralis (Waagen).

PI. ii, figs 8, 9.

1S84-. — Pivdvctus spiralis Waagen, p. 681, pi. Ixvii, fig, 6; pi. Ixviii, fig. 3;
pi. Ixix, figs. 1 1 2, 3.

1916. — Prodnetus spiralis Waagen, Rroili, p. 11, pi. cxvii, figs. 1-5.

1927. — Pradiictus spiralis Waagen, Piener, p. 26.

1928. — Productus spiralis Waagen, Hamlet, p. 16.

Material. — Axis. Miis. F 36513, 36514, 36515, W'yndliam Gap, North-West
Division, Western Australia. Callytharra stage.

Diagnosis . — Shell large, regularly curved through 270°. Hinge-line equal
to greatest width. Ears large and reflexed. Pedicle valve highly inflated with
arched umbonal region, sides parallel and .steep. Ornamentation reticulate on

Permian Productinae and Strophalosiinae of W.A.

19

posterior third of shell, eostate on anterior two-thirds, no development of
folds on trail. Slight convergence of costae in sinus. Ornamentation on ears
partly reticulate, partly costate, costae showing spiral arrangements. Row of
spines along hinge-line and along line of separation of ears and flanks.

Brachial valve flatly concave with strong geniculation. Ornamentation
reticulate on visceral disc, costate on trail. Ears deeply excavated, smooth
or with concentric wrinkles only. No spines. Cardinal process short, trifld.
Well developed adductor muscle impressions. Muscle and brachial impressions
of typical productid pattern.

Description. — Only pedicle valves are known from Western Australia.
These have the following dimensions : —

Description.

1.

2.

3.

Length of hinge-line

88*2

63-0-f

Height

45*6

58-1

48-9

Curvilinear length

134-0

112-0

99-4

The shell is large, as shown by the table, and curved in a regular spiral,
there being no pronounced change in the rate of curvature throughout the
period of shell growth. The umbo, though incurved, is blunt and scarcely
overhanging the hinge-line. The hinge-line is straight and long, and forms
the maximum breadth of the shell, about one quarter of this width being
occupied by the ears alone. These are prominent features of the shell, they
are strongly reflexed with a triangular outline. Their surface makes an
obtuse angle with that of the flanks. A median sinus arises near the umbo
and becomes deeper towards the anterior margin.

The ornamentation is reticulate on the posterior third of the shell, the
rugae and costae being equally developed, thus giving an even pattern. On
the trail the costae become more irregular, this irregularity being due to a
change in disposition of the costae above and below the large spines which
develop sporadically in this part of the shell. No grouping of the costae
into fasciculi is seen in this species. A row of strong spines occurs along
the hinge-line and along the flanks at the line of separation of the ears.
The ornamentation of the ears is one of the characteristic features of the
species. No new costae arise along the hinge-line after the shell ornamenta-
tion has ceased to be reticulate and those already formed become more widely
separated as the shell grows and, following the curvature of the valve, show a
spiral arrangement when viewed from the side.

Comparison with other species. — The spiral ornamentation of the ears
is a distinctive character of this species, and differentiates it quite clearly
from Dictyoclostus caUytharrensis. The visceral cavity is much shalloww in
the latter species, not showdng the arching in the umbonal region which is
seen both in D. spiralis and in D. callytharrensis var. wadei. In both
D. callytharrensis and its variation the costae on the trail are grouped into
fasciculi, a condition not seen in D. spiralis.

For the distinction of D. spiralis from the Indian members of the
spiralis-suhcostatus reference should be made to Waagen (1884).

20

K. L. PREXDERGAST.

LINOPRODUCTUS Chao.

1927. — Chao, Falaeoni. sinica, ser. B, V, fasc. 2, p. 128.

1928. — Chao, ibid, fasc. 3., p. 63.

1931. — Paeckehuann, pp. 75, 206.

Genotype. — Productus cora d'Orbigny, 1842, Vov. Amer, Merid., vol. 3, Pal.,
p. 55, pi. V, figs. 8-10.

Diagnosis. — Thin-shelled Productids with convex pedicle valve; brachial
valve flat or concave in visceral portion, sometimes geniculated anteriorly.
Hinge-line equal to or less than the greatest width. Both valves without
-cardinal area. Pedicle valve evenly convex or sinuate.

Surface ornamentation of fine radiating striae and indistinct concentric
wrinkles; wrinkles more marked on the brachial valve. Scattered spines on
striae present or absent.

Pedicle valve with marginal ridges weak and muscles not strongly
impressed. Median septum in brachial valve. No brachial cones.

Bemarlx'S. — Chao has divided Linoproductids with wavy undulations and
pustulose spines into two groups, one characterised by P- cancriniformis
(Tschernyschew), and the other by P. villiersi d^Orbigny. These groups are
separated by the character of the brachial valve, regularly concave in
P. villiersi and geniculate in P. cancriniformis.

Linoproductus cora (d’Orbigny).

1842. — Product'us corn d’Orbiguy, p. 55, pi. v, figs. 8, 10.

1911. — Producius cora d ’Orbigny, Diener, p. 19, pi. iii, figs. 3-13.

1914.“ProdHcf«.S‘ cora d ’Orbigny, Kozlowski, p. 48, pi. iv, fig. 19 ; pi. v,
fig. 5; pi. vi, figs. 1-10; text-fig. 8.

1916. — Productus cora d’Orbigny, Broili, p. 19, pi. cxv (1), figs. 14-15;
pi. cxvi (2), figs. 1-3.

1927. — Productus cora d’Orbiguy, Chao, p. 132, pi. xiii, figs. 17-18; pi. xiv,
figs. 1-4.

1927. — Productus cora d’Orbigny, Diener, p. 24.

For complete synonymy see Broili (1916) and Diener (1911).

Diagnosis. — Linoproductids of variable size, transversely oval to elon-
gated oval in shape, hinge-line equal to gi'catest width of shell.

Pedicle valve swollen, evenly convex in longitudinal profile, medially
rounded or with broad sinus ; tendency to spread and flatten in fan-like form
anteriorly; flanks sloping steeply from visceral portion. Shell arched over
hinge-line, umbo incurved, slightly overhanging the hinge-line. Ears flat,
their surface perpendicular to flanks of valve. Ornamentation of fine raised
and rounded striae, subparallel or flexuous and irreg^ular; increasing in num-
ber by intercalation of new striae between those pre-existing. Concentric
folds on ears, not crossing visceral disc. Row of spines along hinge margin,
spines present or absent on rest of valve.

Description. — Productus cora was first described by d’Orbigny from
South America, but the incompleteness of his description led to confusion in
the interpretation of the si^ecies for many years. The examination of a
large number of topotypes by Ivozlowski in 1914 showed that there was a
large variation Avithin the species, and his description is here summarised. It
varied in shape from transversely to longitudinally oval; the pedicle valve
might be very SAVollcn in the median region or only moderately so ; the umbo
was sometimes, but hot always, strongly incurved; the pedicle valve Avas

PERillAN PrODUCTINAE AND StROPHALOSHNAE OF W.A.

21

.sometimes impressed by a median sinus, sometimes regularly curved trans-
versely; the striae varied in hexuosity and in the number of spines borne
on the body of the valve. Each of these varied independently of the others,
except possibly that the flexuosity of the striae was correlated with ihe
presence of spines, one or more uniting, a spine arising at the point of union
.and the striae then disassociating.

Interiorly the dendriiic adductor impressions of the pedicle valve are
elongated and parallel to the median line while the diductor impressions are
.short, broad and longitudinally striated, arising at the loAver edge of the
adductors. The brachial valve has a raised median septum, adductors

dendritic on each side of the septum, reniform impressions running laterally
from the base of the adductors, following the lateral and antero-lateral
margins, then returning to the point of origin. Cardinal process trilobed,
median lobs raised above the two laterals. Marginal ridges arising at the
base of the cardinal process and continuing along the hinge to the ears. A
hollow separating the ears from the visceral disc, runs obliquely from the
base of the cardinal process.

Remarks, — Diener in 1911, after examining a collection of Linoproductid
shells from the Fusulina Limestone of Kehsi Mansani suggested that P. coru
d’Orbigny should be enlarged to include those forms previously assigned to
the species P. neffedlevi Verneuil, P, corriigatus McCoy, P. prattenianus
Norwood and Pratten and P. Uneatus Waagen. In this collection were forms
which, Avhile they could not be separated from the type specimens of these
.species, formed a continuous series with the specimens of undoubted P. cora.
Diener’s conclusions that P. cora included a wide variety of forms has since
.been strengthened by Kozlowski's work on the topotypes of P. cora. Koz-
lowski regards P. prattenianus and P. Uneatus as members of the ‘^cora'^
species group. He adds, too, P. ovaius Hall, but thinks that there is a
'detinite, if unimportant, distinction between P. cora and 1\ corragaPus in
that in the latter species the spines arise on any striae at its confluence with
several others whereas in P. cora it is usual for the same stria to bear several
s})ines at different places along its lengtli.

These two authors have each had access to a large lutmber of specimens
and have, each independently of the other, found that P. cora is a very
variable species and includes in itself P. Uneatus, P. prattenianus, P. neffedievi
and P. ovatus. The conclusion of such men, working with good material
must be accepted, and even while these groups may be distinguished by
varietal names, it must be rememl)ered that they all belong to the one species
■group — Prodnctiis cora d’Orbigny.

Australian Bistrihution. — This species has been previously recorded and
figured from Tillighary, N.S.W., by de Koninck. The specimens were, how-
ever, burnt in 1882. Other references to this species in Australian literatiu’e
(Etheridge jun. 1872, P]theridge and Jack 1892, and Etheridge and Dun
1909) are not included in the synonymy of the species as in all the specimens
figured the concentric markings are strong on the pedicle valve, a condition
not showui in any of KozloAvski’s or Diener ’s figures. These S]iecimens* have
not been (examined, and consequently no ojiinion is expressed on the validity
of the identifications.

Productus cora has been recorded from Western Australia by Chapman
:and Glauert (1910, p. 87), It was recorded from the Kimberley District by

90

K. L. Prendergast.

Chapman in 1924 (p. 36), but the specimen has since been re-examined by
Miss Hosking, and re-identified as Streptorhynchus luiluigui (Hosking 1932^
p. 45).

Linoproductus cora var. foordi (Eth. hi).

I’l. iii, figs. 3-5.

1890 . — Froducius temiisiriatus de Verneuil, Foord, p. 151, pi. vii, figs. 4, 4a..

1903. — P. te7iuisfriatus de Verneuil, var.. foordi Etheridge, jun., p. 19, pi. i,,
figs. 3-4; pi. iii, fig. 22.

1907. — P. tenuistriatus var. foordi Etheridge, jun., p. 30, pi. ix, figs. 4-6.

1910. — P. tenuistriatus var. foordi Eth. fiL, Glauert, p. 87.

1931. — P. tenuistriatus var. foordi Eth. fil., Hosking, pp. 8, 22.

Specimens. — G.S.W.A. 1/4683, south bank of Wooramel River below Cally-
tharra Spring, Callytharra Limestone.

G.S.W.A. 5708b., Fossil Cliff, Irwin River.

Aus. Mus. F 36247, 36251-36255 (inch), Gascoyne River, near Winnemia^
North-West Division. Byro Stage.

Aus. Mus. F 9027, Wyndham River, North-West Division. Byro Stage

Aus. Mus. F 36287, Gascoyne River, North-West Division. Byro Stage.

U.W.A. 10822, Fossil Cliff, Irwin River. Fossil Cliff horizon.

Diagnosis . — Linoproductids with elongate oval outline, hinge-line equal
to gi’eatest -width of shell.

Pedicle valve without median sinus. Visceral region very swollen, arch-
ing over hinge-line, umbo slightly overhanging hinge-line. Ears small and
hat. Valve regularly curved in longitudinal prohle. Striae hne, irregular,,
increasing by intercalation along a definite zone. Concentric wrinkles weak,,
not crossing visceral disc. Groups of spines at cardinal angles and one row
at low angle to hinge-line.

Brachial valve concave, ornamentation as in P. cora d’Orbigny.

Description . — The pedicle valve has an elongated oval outline in the
larger specimens. It is gibbous in the visceral region falling rapidly at the
sides to small, flat ears, and truncated posteriorly by a small umbo over-
hanging the hinge-line. Anteriorly there is a decrease in the curvature so
that the shell is elongated rather than globular. A row of spines along the-
hinge, at about 10° to the cardinal margin terminates in a group of from
4-6 spines on each ear. No spines occur on the rest of the shell. The longi-
tudinal striae are fine (20-40 measured over a breadth of 20 mm. at a dis-
tance of 10 mm. from the umbo), and vary in irregularity. On some speci-
mens they may run almost parallel for the length of the shell, but more
usually two or more unite and re-divide later, having an irregular course.
The striae increase by intercalation and this takes place in most specimens,,
at about the same stage of growth. The amount of increase, too, varies, one
or two striae being intercalated between a pair of primaries. Folds on the-
ears are not seen on the majority of specimens and where they are present
never cross the visceral part of the shell.

The brachial valve is concave, the ornamentation as in the pedicle valve,,
but crossed by concentric wrinkles. The valve is devoid of spines.

Permian Productinae and Strophalosiinae of W.A.

23

The sizes of the specimens (pedicle valves) are shown by the following
table : —

1 .

2.

3.

4.

5.

6.

Height

29-1-f

22-8-h

31-0 +

23-7 +

23-04-

14-1

Maximum width

29-4 +

25- 1-h

27-0-f

27-04-

24-0 +

13-3

Length of hinge-
line

22-9 +

20'2-y

19-84-

25-0

18-6 +

13-3

Curvilinear length

50-0-1-

39-3-h

56-0 +

41-14-

35-5-f

21-8

No specimens examined have both valves in position, it is thus impossible to
estimate the thickness of the shell.

The width of the hinge-line as shown by these figures is less than the
maximum width of the shell. The ears in these specimens are very fragile
.and readily break off; the only complete specimen is a small one (No. 6).

The internal structures are as described for the species P. cora^ the
muscles being inserted in definite fossae.

Comparison with other forms. — These specimens have been referred for
many years to Prodiictus temiistriatus Verneuil, although distinguished as a
variety. Etheridge (1903) separated them as a variety giving as his reasons
the much coarser and more irregular ornamentation (in P. temiistriatus there
are 50-60 striae in 20 mm. at a distance of 20 mm. from the umbo). They
differ from that species, too, in the width of the hinge-line, the arching of
the i)edicle valve and the regular anterior elongation, P. tenuistriatus being
irregularly produced anteriorly. Etheridge has already said that this form
belonged to the ^^cora’^ group (1907, p. 30). Specimens from Timor (Broili
1916, pi. cxv., figs. 15-16; pi. cxvi., figs. 1-3) probably belong to this variety
‘Oi P. eora.

This variety is not distinct from P. cora d’Orbigny. It is a variety
within that species as can be seen by a comparison of KozlowskPs figures
(pi. vi., figs. 7a, b) and figures accompanying this paper, but a variety which
has become stabilised in this area. The limits of variation are narrower than
in those from South America. The shell shape is constant as is the convexity
■of the pedicle vaNe, it is not sinuated and does not carry spines on the main
part of the shell. The varietal name is retained for these specimens until
the complete range of Productus cora d’Orbigny is seen in this country.

Linoproductus cancriniformis (Tschernyschew).

1889. — P. cancriniformis Tschernyschew, p. 373, p]. vii, figs. 32, 33.

1897. — P. cancriniformis Tsehern. Diener, vol. 1, Pt. Ill, p. 25, pi. vi, figs,
fia-b, 7a-d.

1897. — P. cancriniformis Tsehern. Diener, vol. 1, Pt. TV, p. 31, pi. 1, figs.

1907. — ^‘P, nndatus^^ Def ranee, Eth. fil., p. 30.

1916.- — P. cancriniformis Tsehern., Broili, p. 13, pi. cxvi, fig. 6.

1918. — ‘^P. hellns’^ Eth. fil., p. 254, pi. xxxix, fig. 4-5; pi. xl, fig. 6.

1928. — L. cancriniformis (Tsehern.), Chao, p. 65, pi. v, figs. 8-9.

1934. — L. cancriniformis (Tsehern.), Prendergast, p. 14, pi. ii, figs. 7-12.

1936. — L. ca7ieriniformis (Tsehern.), Douglas, p. 30, pi. iv, fig. 3.

24

K. L. Prexdergast.

MatericK — Aus. Miis. F 38453-38457, 9069, Minilya Biver, North-West
Division. Wandagee Stage.

Aus. Mus. P 37580, Vi mile west of shale outcrop. Northern bank. Same-
locality and horizon.

Aus. Mils. F 37576, % mile cast of shale outcrop. Northern bank. Same
locality and horizon.

Aus. Mus. F 16729, 10749, Irwin River District Fossil Cliff horizon.

Aus. Mus. F 37565, ^4 mile east of fault, south bank of Minilya River,
North-West Division. Wandagee Stage.

Diagnosis . — Linoproductids with non-sinuate pedicle valve, hinge-line'
equal to greatest width of shell, umbo overhanging hinge-line; ears distinct
but not prominent. Shell ornamentation of fine striae, becoming swollen
at regular intervals to give anteriorly directed spines, quincuneially arranged.
Row of spines along cardinal margin. Visceral disc crossed by concentric
wrinkles; size and extent of wrinkles variable.

Brachial valve flat over visceral disc becoming strongly geniculated'
anteriorly. Short median septum. Ornamentation as on pedicle valve.

Internal features not known.

Description . — For description and discussion of this species see Prender-
gast, 1934, p. 14.

Linoproductus cancriniformis var. lyoni var. nov.

PI. iii, figs. 1-2.

Jfolofypc-. — Aus. Mus. F 36530, 10 chiis. north-west of Giiarrca Pool near-
Winning Statin, North-West Division. Lyons Stage.

Topotypes. — Aus. Mus. P 36533, 36535, 36537, 36538, 36540. Same locality
and horizon.

Diagnosis . — Shell medium to large, semi-circular outline. Pedicle valve
evenly convex, not swollen, without median sinus. Hinge-line straight, equals,
greatest width of shell. Umbo strong, not incurved, not overhanging hinge-
line. Ears large, fiat, not clearly differentiated from body of shell. Orna-
mentation as in Linoproductus caneriniformis Tschern.

Brachial valve and internal structures of pedicle valve not known.

Description.

Dimensions.

36530.

36531.

36532.

Length of hinge-line

43-7

34-5

22*7

Height

32-6

28-0

21-5

Curvilinear length of pedicle valve

41-0

25*8

34-2

The pedicle valve has a semi-circular outline with the greatest Avidth at
the hinge-line. The longitudinal profile of the shell is a regular cuiwe be-
coming geniculate near the front; transversely the curve is regular, without
any median flattening. The ears are large Avith alar angles right angles,
they are not distinct from the body of the shell. The shell surface is
covered Avith fine radial striae (10-13 in 10 mm. measured at a distance of
20 mm. from the umbo) bearing swollen elongated spine-bases arranged in;

Permian Productinae and Strophalosiinae of W.A.

2d

<tuincimx. The striae increase in number by division of primaries. Con-
•centric wrinkles are present on the ears but these rarely extend onto the body
'Of the shell. A row of large erect spines is found along the cardinal margin.

Comparison with other species. — Only pedicle valves are known of this
variety. These, however, are such a distinctive shape that it has been thought
best to distinguish them temporarily as a variety of Linoproductus can-
■criniformis (Tschern.). The ornamentation of the variety is similar to that
■of Tschernschew^s species with the same fine radial striae and elongated
pustules. They differ in the much less convex pedicle valve and in the semi-
circular outline with the greatest width at the hinge-line.

The variety shows also a marked resemblance to Productus asperidiis
Waagen (1884 p. 693 pi. Ixxix. figs. 3-6) in outline and type of ornamenta-
tion. The spines in that species are erect and not elongated pustules seen in
Linoproductus cancriniform'.s.

Productus koninchianns de Vern. (Chao 1928, p. 63 pi. v, figs., 4-7) has
also some features in common with these shells. It has, how^ever, a much
more inflated pedicle valve though several figures of smaller specimens
(Chao 1928, pi. v. fig. 6) have a fiat pedicle valve. The Western Australian
;specimens are always larger than these smaller shells and never have an
inflated and incurved umbo. Productus tumidus Waagen (183-1, p. 708,
pi. Ixxx. figs. 1-3) is also an inflated form and bears erect spines.

WAAGENOCONCHA Chao.

1027. — Chao, Y. T. Palaeoirtt. sinlca, S?r. B., V, fasc. 2, p. 8o.

Diagnosis. — Thin-shelled productids with convex sinuate pedicle valve,
brachial valve flat or concave. Surface marked by quincuncially arranged
tubercular spines in the young and adult stages, becoming smaller and more
-closely packed together in old age.

Gonctype. — Productus humholdti d’Orbigny, 1842, Voy. Amer. Merid., Vol.
3, Pt. 4, p. 54, figs. 4-7.

Waagenoconcha imperfecta Prendergast.

PI. iii, figs. 7-9.

1924 . — ‘‘Productus suhquadraUis^ ’ Morris, Chapman, p. 36.

1934 . — Waagenoconcha imperfecta Prendergast, p. 15, pi. iv, figs. 1-3.

Holotype. — U.W.A. 3044, Luiluigui Station, Kimberley Division. Liverynga
Series.

Allotypes. — U.W.A. 2768, 2775, same locality and horizon.

Topc-t-ypes. — U.W.A. A 444, Lniluigni HomciLttet d, Kimberley Division.
Liverynga Series.

Other Materifd, — U.W.A. 20457, Limestone at base of Mt. Hardman, Kim-
l)erley Division. Liverynga Series.

U.W.A. 20454, north flank of Mt. Cedric, Kimberley Division. Liverynga
iSeries.

-Shell transversely oval in outline, hinge-line less than greatest
width of shell. Ornamentation on both ^alves of coarsely spaced quincun-
cfally arranged spine bases with fine forward projecting spines. Shell with
well marked growth lines.

Pedicle valve evenly convex, sinuate, umbo prominent, produced beyond
hinge-line.

Brachial valve fiat to slightly concave vdth a shallow median fold.
Alargins not geniculate.

26

K. L. Prendergast.

Description . — See Prendergast 1934, p. 15.

Demarks . — In the earlier specimens of this species examined it was-
noted that the specimens did not agree with the generic description since the-
spines were less crowded anteriorly. Specimens in a better state of preser-
vation have since been obtained and these show that the inclusion of the
species in the genus Waage^ioconcha is quite justified. In these specimens the
spines are crowded together on concentric wrinkles on the older parts of the
shell. The absence of this feature on the earlier specimens Avas undoubtedly
due to exfoliation of the outer layers of the shell leaving only coarser pustules
more widely spaced.

When this species was originally described no copy of Broili^s (1916)-
work on the Permian Brachiopoda of Timor was available and a new species
was founded to contain specimens which belong to Productus \caageni Roth-
pletz as figured by Broili. Comparison of the Western Australian specimens
with Broili’s figured specimens and with others from Timor (Basleo, Neoet-
pantoekak, Toenion Eno, Timor. B.M. B. 98397, 98985, 98648) leaves no
aoubt on this point. The Western Australian name is retained because it is'
doubtful if P. waageni Broili is identical with P. waageni Rothpletz.

Rothpletz’s figures of P. waageni (1892, pi. x. fig. 19) show a small shell
transversely oval in outline, ornamented Avith fine closely set spine bases. It
shoAvs no groAvth lines and has a very Aveak sinus in the pedicle valve. But
Rothpletz compares his specimens Avith P. lmmhold,ti as figured by Waagen
(1884, pi. Ixxvi. figs 1-3), a rather surprising comparison as most of the
specimens figured by Waagen have Avell-marked concentric ornamentation.
Indeed Rothpletz’s figures (pi. x. fig. 20) of P. ahiclii are more like P. kum-
boldti AVaagen. If P. waageni Rothpletz is identical A\fith P. u'aageni Broili.
then the absence of growth lines on Rothpletz's type is due to AA’eathering or
to individual variation. Without inspection of the type specimen of P..
waageni Rothpletz, this species must be regarded as Avithout growth lines.

Waagenoconcha vagans Reed.

PI. iii, fig. 6.

19.31. — Waagenoconcha vagans Reed, p. 9, pi. ii, fig. 2.

Material . — One brachial valve, IT.AV.A. 20459, north flank of Mt. Cedric,,
Kimberley Dhfision. LiA'erynga Series.

Diagnosis . — Shell trapezoidal to almost square in outline, hinge-line less
than the greatest Avidth of shell.

Pedicle valve sinuate, umbo SAA^ollen and overhanging hinge-line, ears
fiat. Ornamentation of small spine bases quincuncially arranged, pits
occurring betAveen spine bases.

Brachial valve fiat Avith shalloAv median folds, margins geniculated.

Description . — This s])ecies is knoAvn in AVestern Australia only as an-
external cast of a brachial valve.

The shell is trapezoidal in outline, the greatest Avidth at the anterior-
margin, this and the hinge-line being the parallel sides. The hinge-line is
short and ecpial in length to half the maximum Avidth of the shell. The fiat
visceral disc is divided medially by a shalloAv fold, the fold AA-idening an-
teriorly. Laterally and anteriorly the margins of the valve are geniculated to
form a fringe. The surface of the shell is ornamented by numerous holloAvs
and spine bases quincuncially arranged (18 spine bases in 10 mm. 20 mm.
from umbo), resembling the ornamentation of IT. imperfecta mihi. Numerous^
groAvth lines cross the surface of the valve.

Permian Productinae and Strophalosiinae of W.A.

27

From its impression the umbo of the pedicle valve seem to be swollen
■and overhanging the hinge-line.

The brachial valve has a length of 29 mm., with a maximum width of
-38 mm.; the length of the hinge-line is 24 mm.

Remarks , — This species was first described by Reed from the middle
Productus Limestone at Kumaranwali in the Warcha Valley of the Western
Salt Range. It is characterised by the ornamentation in conjunction with the
geniculation of the brachial valve.

TAENIOTHAERUS Whitehoiise.

1928. — Whitchouse, }iep. Aust. Ass. Adv. Sci., vol. xviii, pp. 281, 282.

Genotype. — Productus svA) quadrat us Morris, 1845, in Strzelecki, Physical
description of N.S.W. and Van Diemen's Land, p. 284.

Diagnosis . — Adult shell large, general outline subquadrate to elongate
■oval; pedicle valve evenly convex with median sinus; brachial valve slightly
■convex to flat with upturned margins. Pedicle valve sometimes with trian-
gular concave area; delthyrium, where present, partially filled by triangular
■extension of cardinal margin of brachial valve.

Ornamentation of coarse elongated spine bases possibly giving rise to
long spines; spine bases not continuous into costae. Irregular concentric
wrinkles or lamellae widely spaced over whole shell.

Muscle impressions as in Aulosteges and Productus. Brachial valve with
median septum. Cardinal process large, varying angle of inclination to ear-
‘dinal margin.

Remarks . — For description of this genus see Prendergast, Proc, Roy.
■Soc. Tasmania.*

Taeniothaerus subquadratus (Monas) .

PI. iv, figs. 1-6.

1845. — Productus suhquadratus Morris in Strzelecki, p. 284.

l^^2.~Productus svhquadratus Morris, Etheridge and Jack, u. 282 ul.
xxxviii, figs. 7-10; pi. xl, fig. 5. '

1907. — P. subquadratus Morris, Etheridge, jnn., p. 21, pi, iii, figs. 6 and 7
(in the plate fig. 7 is labelled as fig. 1).

1907.— P. snhquadratus Morris (?), Etheridge, jun., p. 30, pi. vii, figs. 2, 4;
pi. viii, fig. 1.

1909. — P. suhquodratus Morris, Etheridge and Dun, p. 300, pi. xli, figs. 1-5.

1910. — P. subquadratus Morris, Glauert, p. 87.

1914. — Aulosteges baracoodensis Eth. fil., Etheridge, jun., p. 33, pi. iv, figs.

1931. — P. mibquadratus Morris, Hosking, pp. 8, 22.

1933. — P. subquadratus Morris, Hosking, p. 36.

1933. — Aulosteges cf. A. spinosus Hosking, p. 36.

1935. — P. subquadraPus Morris, Prendergast, p. 17, pi. iv, fig. 4.

Jlolotype. — B.M. 91171, Mt. Wellington and Mt. Dromedary, Tasmania.

Other Material— U. W.A. 4768, 12396, 20450, 21247, Fossil Cliff, Irwin River.
Fossil Cliff horizon.

U.W.A. 10931, two miles north of Ballythanna Hill, Wooramel River Dis-
trict. Byro Sandstone.

U.W.A. 20447, below Coolkilya Pool, Minilya River, North-West Division.
Wandagee series.

G.S.W.A. 10930, Mt. Marniion Kimberley Division. Exact horizon not known,
G.S.W.A. 10929, locality and horizon not known.

G.S.W.A. 1/4655, locality and horizon not known.

* In January,
of Tasmania on this
appeared. — E. deC.

1941, Dr. Prendergast had completed a paper for the Royal Society
subject, and was awaiting the illustration^i, but the paper has not vet
Clarke, 22/3/43.

28

K. L. Prendergast.

Diagnosis . — Shell large, subquadrate to oval in outline. Greatest width-
near anterior margin. Pedicle valve strongly and evenly convex with median
sinus, with or without cardinal area. Brachial valve slightly convex to con-
cave with upturned margins.

Ornamentation essentially spinose. Spine bases elongated posteriorly..
Concentric lamellae or wrinkles crossing shell surface.

Internal features as in Prodactus s.l. Cardinal process vertical to in-
clined almost to horizontality. Median septum in brachial valve.

Description . — This species has been described and discussed in Prender-
gast {Proc. Pog. Soc. Tas.)*, but some special features of the Western
Australian shells are added here.

The pedicle valve is strongly convex with steep flanks and indented by a;
median sinus, which flattens out anteriorly so that the margin may be entire. -
The area is narrow in some specimens but may become wide. It is divided by
a definite delthyrium which is partially filled by a triangular projection of
the brachial valve. The adductor impressions are dendritic and placed high
in the valve, the longitudinally striated diductors arise at the sides of the
adductors and are produced laterally and anteriorly.

The brachial valve is flattish with a slightly convex region near the
umbo; it is upturned at the margins. The cardinal process is strong and
trilobcd and varies in its inclination to the plane of the valve from being
eoplanar to almost perpendicular, the process in this case assuming a horizon-
tal position. The median sei^tum arises at the base of cardinal process and
continues forward for about two-thirds of the length of the valve. The adduc-
tor muscle impressions are dendritic and enclosed in a heart-shaped elevation
whose apex is at the base of the cardinal process. From the base of this eleva-
tion arise the brachial impressions, these running parallel to the hinge-line’

almost to the lateral bordei’s then turning in a circular path and returning to
their points of origin parallel to their outward path. At the base of the
cardinal process arise two laterally directed ridges running parallel to the
hinge-line but dying out before reaching the ears.

The ornamentation of the shell varies within rather wide limits. It is
essentially spinose but the intensity of the spinosity varies from locality to
locality. In the Irwin shells and those from the North-West Division the
spines are very fine and closely set on concentric lamellae, in those from the
Kimberley Division they are much thicker and fewer in number.

* See footnote p. 27.

Permian Produotinae and Strophalosiinae op W.A.

29

Remarks. — From the Fossil Cliff Beds, Irwin River, are found shells
which possess an elongated narrow area. Those specimens from the Byro
sandstone of the Wooramel River have a distinct though not large area, while
a specimen from the Kennedy sandstone (North-West Division) has been
identified as Aidosteges ingens Hosking, although it differs from the Byro
specimens only in the greater development of the area and the horizontality
of the cardinal process. The stratigraphical relationships of these beds are
given by Raggatt (1936) and Rudd and Dee (1932, MSS. unpublished).

N.W. Division and

W^ooramel River. Irwin River.

Gascoyne Series
5. Wandagee
4. Kennedy
3. Byro
2. Wooramel

1. Callytharra = Fossil Cliff Beds,

The specimens of these species may, then, be arranged in a time series,
in which the area increases in size as the fossil-bearing rock is 3 munger in
age. The cardinal process also changes gradually through this series. In the
typical Productus (e.g. Dictyoclostus semireticulatiis (Martin)) the cardinal
process is erect and in specimens with small areas this statement also holds
for Taeniothaerus subquadraius (Morris). In Aulosteges haracoodensis Eth.
fll., however, as in all Aulosteges the cardinal process has become bent through
a right angle. This change, from vertical to horizontal, takes place gradually
and keeps pace with the growth of the cardinal area. The suggestion is there-
fore made that Taeniothaerus suhquadratus (Morris) is a transitional form
connecting an unknown Rroductus with Aidosteges haracoodensis Eth. fil.

KEOTOVIA Fredericks.

1928. — Fredericks, G., Bull. Com. Geol. St. Petersh., 46, 1927, p. 790.

Genotype. — P. spindulosus J. Sowerhy, 1814, Min. Conch., Vol. 1, pi. Ixviii,
fig. 3.

Diagnosis . — Small thin-shelled product!. Pedicle valve convex, brachial
valve concave to geniculate, closely following curve of pedicle valve. Orna-
mentation of fine spines in irregular quincuncial arrangement covering whole
shell surface; spine bases elongated in anterior part of shell. With or with-
out weak concentric wrinkles, median sinus only slightly developed or absent.

Internal structures unknown.

Comparison ivith other genera. — This genus differs from Fustida Thomas
in the absence of marked concentric wrinkles and in the nature of the visceral
cavity. In Pustida the brachial valve is flatfish, forming a large visceral
cavity, whereas in Krotovia the brachial valve follows the pedicle valve so
closely that the visceral cavity is thin. Schuchert and Le Vene consider that
this is not sufficient distinction as they say (1929, p. 72) of Krotovia “appears
to equal Pustula Thomas.'^ The shape of the body cavity is, however, a
feature of diagnostic importance, and I have, therefore, adopted Frederick’s
genus.

Frederick’s diagnosis of Krotovia (p. 790) states: “The shell has an
oval configuration and a plain surface. The ornamentation consists of spines
only,” and in the Russian text (p. 779) “Sometimes irregular ribs and folds

30

K. L, PRENDERGAST.

develop which are connected with the spines.’' Krotovia then includes speci-
mens with incipient plications thus overlapping with Avonia Thomas.
Paeckelmann (1931, p. 77) has emphasised this fact, and distinguished
Avo7iia s.str. and Krotovia as subgenera of Avonia Thomas, the forms to be
separated by the presence {Avo^iia) or absence {Krotovia) of costae. The
distinction is thus purely arbitrary, the two genera intergrading.

1932 . — Pusiula senticom Hosking, p. 47, pi. iii, figs. 2-3.

Syntypes. — G.S.W.A. 1/4970 (a), from Creek 1 mile west of Callytharra
Springs, Wooramel River. Callytharra Limestone

Biag^iosis. — Thin-shelled form, outline semi-circular to subquadrate,
hinge-line less than maximum breadth. Pedicle valve gently convex, non-
sinuate, auriculate, umbo pointed and incurved. Brachial valve strongly
concave; visceral cavity of Productus typici ^ type. Ornamentation on both
valves of erect spines in irregular quincunx and fine concentric lamellae.

Internal features of pedicle valve unknown, brachial valve with short,
broad cardinal process continuing into short median septum. :

Description. — See Hosking, 1932, pp. 47-49. i

Pemarks. — The type specimens of this species have a convex pedicle i
valve with a concave brachial valve, the visceral cavity between the tw^o
being very thin. Also, as Miss Hosking says in her description (p. 48) “the
lack of any marked concentric ornamentation is one of the chief features of
the species.” The specimens must, then, be referred to Krotovia rather than
to Pustula.

1932 . — Pustula micracauiha Hosking, p. 49, pi. iv, figs. 4a, b.

Syntypes: — G.S.W.A. 1/4970 (b), from Creek Vj mile west of Callytharra
Spring, Wooramel River. Callytharra Limestone.

Diagnosis. — Small, thin-shelled form. Shell outline semi-circular to sub-
quadrate, hinge-line slightly less than greatest shell breadth. Pedicle valve
geniculate, non-si nuate; umbo small, not overhanging hinge-line. Brachial
valve deeply concave and geniculate, closely following contour of pedicle
valve ; visceral cavity of Productus typici ^ type. Ornamentation on pedicle
valve of recumbent spines in irregular quincunx with concentric folds pos-
teriorly and on ears.

Short median septum in brachial valve. Other internal features not
known.

Description. — See Hosking, 1932, pp. 49-50.

Remarks. — From an examination of the type specimens there can be no
doubt that this species should be referred to Krotovia. The visceral cavity is
thin, the concentric wrinkles are not pronounced and the nearest British form
is P.spiniilosus J. Sow., the genotype of Krotovia.

1814 . — Productus spinulosus J. Sowerby, p. 155, pi. Ixviii, fig. 3. }

1860 . — Productus spinulosus J. Sowerby, Davidson, p. 182, pl. ii, figs. 23, 24. [

Krotovia senticosa (Hosking) .

Krotovia micracantha (Hosking).

Krotovia spinulosa (J. Sowerby).
PI. iv, figs. 11-13.

I

Permian Productinae and Strophalosiinae or W.A.

31

1861. — Froductus spinulosus J. Sowerby, Davidson, vol. ii, part 5, No. 4,
p. 175, pi. xxxiv, figs. 18-21.

1914. — Pusttila spmulosa J. Sowerby, Thomas, p. 314, pi. xviii, figs. 7-9;
pi. xix, figs. 7, 8.

1930. — Krotovia spinulosa (Martin), Muir-Wood, p. 106. (There is no Pro-
ductus spinulosus Martin — this is a mistake in author of species) .

Jloloiype. — Present location unknown, listed as Royal Scottish Museum,
Edinburgh, Pleming Collection. Carboniferous Limestone, West Lothian, Lin-
lithgowshire.

Material. — Aus. Mus. E 38442, 38443, 38446, Wandagee Station, Minilya
River, North-West Division. Wandagee Stage.

Aus. Mus. F 37563, 37564, 37578, 37582, bank of Minilya River, North-West
Division. Wandagee Stage.

Aus. Mus. F 37725.

Aus. Mus. F 37579, ni. west of shale outcrop, northern bank of Minilya
River, North-West Division. Wandagee Stage.

Diagnosis. — Shell small. Outline oval, hinge-line less than maximum

width. Pedicle valve convex, non-sinuate, umbo small, pointed and incurved,
overhanging hinge-line; ears small and flat. Brachial valve evenly concave.
Visceral cavity of Productus typici jS type. Ornamentation on both valves
of fine oblique spines, closely spaced, with quincuncial arrangement. De-
velopment of incipient costae below spines near anterior margin.

Internal features not known.

Dimensions.

Description.

Type.

F 38443

F 38446

Height

16.2

16.5

Length of hinge line

10.5+

12.5

Maximum width

11.7+

17.3

Curvilinear length of pedicle valve

29.4

27.4

The shells of this species, as may be

seen from

the table.

are small.

Usually transversely oval, they have a straight hinge-line shorter in length
than the maximum width of the shell.

The pedicle valve is evenly convex, without a median sinus. The shell
has its maximum convexity behind the umbo; from this point the sides drop
steeply to fiat ears; anteriorly the slope is more gradual. The alar angles are
obtuse. The umbo is small, pointed and overhanging the hinge-line. The
surface of the valve is ornamented by small closely-set spine bases (10-12 in
10 mm. at a distance of 10 mm. from the irnibo). These give rise to slender
oblique spines on the body of the shell; they have not been observed on the
ears, but a row of erect spines separates the ears from the body of the shell.
On the anterior and lateral slopes small ripples ( ? incipient plications) are
seen anterior to the spine bases.

The brachial valve is regularly concave with the maximum concavity
just beneath the umbo. The ears are flat and in contact with those of the
pedicle valve. The ornamentation is the same as that seen on the pedicle
valve, but no spines are preserved on the brachial valves. A few growth lines,
cross the shell.

Comparison with other species. — Productus opuntia Waagen resembles
this species in many particulars. Both have the same outline, but may be dis-
tinguished by the location of the maximum width which in P. opuntia is the
length of the hinge-line. The spine bases of P. opuntia are not so crowded as
those of P. spinulosus and generally are more robust, having a distinct an-
terior prolongation and are also more nodular.

K. L. Prexdergast.

:v2

Girty (1908, p. 264) compares his species Productus signatiis with
P. opuntia. The fig’ures are not clear, but give an impression of an ornamen-
tation of Cancrinella rather than of Krotovia or \Pustula. The spine bases
are heavy and elongated, and Girty himself says that on some of the casts
irregular lirae can be seen between the spine bases. The species is distinct
from K. spinulosa.

The characters distinguishing the other species of this genus, Krotovia
micracantha and Krotovia senticosa, from this species are dealt with in the
description of those species.

AULOSTEGES llelnierseii.

1847. — Ilelinerscn. Leonhard & Broun ’s Jahrbuch fiir Mineralogie p. 331,
text-figure.

Genotype. — Anlos1< ge,^ variabiUs Helmersen = Orthis wangenheimi (Ver-
neuil). Helmersen 1847, Leonhard and Broun ’s Jahrbuch fur Mineralogie, p.
331, with text-figure.

Diagnosis. — Medium to large shells, ovoid to quadrangular or triangular
in outline. Concavo-convex or platy-convex dorso-ventrally. Ornamentation
essentially spinose with development of radial lirae or concentric wrinkles.

Pedicle valve convex with well-developed area and pseudo-deltidium.
Umbo ])ointed, possibly deformed or with cicatrix of attachment. Edentu-
lous. Shell with median sinus.

Brachial valve flat or concave, without area or area narrow and linear.
Cardinal margin produced into triangular extension medially. Median septum
separating dendritic muscle impressions. Pattern of brachial impressions
productiform. Cardinal process large, inclined or horizontal.

Description. — The shells of this genus are usually massive although
there is a great variation within the genus. They vary in shape from oval
to quadrangular and triangular, the hinge-line being less than the greatest
width of the shell. Neither the i^edicle nor the brachial valve is produced
into distinct ears.

The pedicle valve is regularly convex or geniculate. Ti’ansversely it is
depressed by a median sinus which commences some distance behind the
umbo, is strongest in the centre of the shell and flattens out towards the
anterior margin. The area is triangular and usually well-developed, it is
marked by vertical or longitudinal striations and may he flat or concave;
the concavity may not be regular and a sudden change in curvature gives
the recurved area seen in A. s})inosus, Hosking. A narrow ]'>seudo-delticl!uiu,
triangular or parallel-sided, divides the area. In some specimens, for instance
Aulosteyes ivangenheimi (Verneuil), small s])ines are present on the pseudo-
deltidium. These are not seen in all other species, but their occasional oc-
currence suggests that their absence is secondary, due to removal. The umbo
is sharp and pointed, and usually erect. AVhere, howevei’, the umbo has been
deformed by attachment, it may be twisted to one side or produced across
the area {see, for example, Aulosteges haracoodensis Eth. til., 1914, pi. iv,

fig. 13). The surface of the valve is ornamented by spines, erect and/or

adherent, and may be concentrically wrinkled on the visceral disc. Radial

lirae are seen in some of the species, e.g., A. ivolfcampensis King and A.

medlicottianus Waagen, while stronger radial markings ornament the trail in
A. tibeticus Diener.

Permian Produotinae and Strophalosiinae of W.A.

33

The brachial valve of the genotype A. wangenlieimi (Verneuil) is slightly
convex over the visceral disc, becoming concave with geniculated margins.
This valve has an operculiform character throughout this genus, serving as a
fiattish or slightly concave lid for the visceral cavity. The ornamentation is
as in the pedicle valve.

A description of the internal features of a brachial valve of A. wange^i-
heimi (B.M. BB 32-79) will illustrate the characteristic features of the valve.

The cardinal process is large and trilobate, it is inserted perpendicular
to the hinge-line so that it extends horizontally, not vertically as in the Pro-
ductids. The roots of the cardinal process extend posteriorly, and some part
of them now lies behind the hinge-line. This is protected externally by a
triangular extension of the brachial valve which fills the delthyrium of the
pedicle valve. This structure of the cardinal process is characteristic of
the genus.

At its base the cardinal process is continued into two thickened ridges
which run parallel to the hinge-line, leaving, posterior to them, a rectangular
platform of which the surface of the valve forms the back and the thickened
ridge the floor. The development of these ridges varies within the one species
even under the same environmental conditions; one of the specimens exam-
ined (B.M. BB 3278) had them enormously thickened (see Figure 4), while
in another they were but feebly developed with little laterial expansion. In
the thickened specimen two ridges, directed towards the centre of the shell,
arise at the base of the cardinal process and unite in the middle line to con-
tinue forward as the median septum. These two ridges and the lobes of the
cardinal process form the edges of a diamond-shaped depression which
separates the cardinal process from the median septum. With less-developed
ridges the anterior sides of the depression become obscured.

Figure 4.

Interior of Bracliial Valve of Aulosteges wangenheimi (Verneuil). Permian.
Mt. Grebeni, near Orenburg, Russia. B.M. BB 3279 (X3).

34

K. L. Prexdergast.

The dendritic muscde impressions are situated in sub-triangular depres-
sions on each side of the median septum. At the base of each, the brachial
impression arises, runs parallel to the hinge-line almost to the lateral margin,,
then turns anteriorly to form an open loop. On no specimen have the im-
pressions continued to the median septum. In the area circumscribed by the
brachial impressions the inner shell surface is smooth and shiny, elsewhere
it is finely iDitted.

Figure 5.

Cardinal Process of Aulosteges wangenheimi (Verneuil). Permian. Mt.
Grebeni, near Orenburg, Russia. B.M. BB 3278. (a) vertical, (b) lateral

(c) horizontal (X3).

Coynimrison with other genera. — The genus Aulosteges is distinct from
members of the Strophalosiinae in the possession of a high reclined area,
poor articulation, a cardinal process orientated perpendicular to the commis-
sural plane of the valves, the triangular extension of the cardinal margin of
the brachial valve and the dendritic adductor impressions in the brachial
valve.

The high area of Aulosteges and the horizontal inclination of the cardinal
process serve as distinguishing features from the Productus spinosi group.

In view of the prevailing opinion of Russian palaeontologists that
Aulosteges is a synonym of StropJialosia^ the differences between the two
genera will be considered more fully. Netschajew (1911, p. 144) states that,
the only significant feature distinguishing the two is the presence of deltidial
spines in Aulosteges. Such a statement completely ignores the differences in
the internal structure of the brachial valve. It must be admitted, however,
that the internal features of the Russian species Strophalosia horrescens^
S. fragilis and 8 . gigas are very similar to those of Aulosteges. The muscle
impressions are dendritic, the cardinal process inclined towards the horizontal
and the pattern of the brachial impressions productoid. It is here that the
error lies, for those species mentioned above do not belong to the genus
Strophalosia^ they are Aulosteges,

Licharew (1937, p. 128), while stressing the synonymity of Aulosteges
and Strophalosia, recognised that the Russian “ Strophaloskd^ did not come
within the accepted conception of that genus. He compromised by suggesting

Permian Productinae and Stropralosiinae of W.A.

35

that the S. horrescens group should be distinguished as Aulosteges, a sub-
genus of Stroplialosia. But why a sub-genus? Strophalosia is distinguished
by non-dendritic muscle impressions, a smallish cardinal process, a perclined
area of moderate height in the pedicle valve and well-developed dentition.
"What features, therefore, have Aulosteges and Strophalosia in common that
would induce us to consider one as a sub-division of the other? The posses-
sion of a concavo-convex shell with an area on the pedicle valve and usually
a spinose ornamentation is surely insufficient evidence for this grouping when
so many points of difference are considered.

Fredericks has a more positive reason for distinguishing between the
two genera. He says (1931, p. 209) rhat both genera are characterised by
the presence of an apical apparatus representing a reduced spondylium.
The apical plates have become delthyrial ridges and the secondary septum a
median ridge. These elements, Fredericks says, retain their connection at
the apex of the umbo. He publishes no diagrams in support of this
hypothesis, which rests, therefore, solely on his interpretation of these struc-
tures — structures which would be preserved in only those specimens where
no deformation of the umbo had occurred. The presence of such a structure
would not do more than suggest a similar origin for both genera, a view
which has not been disputed, and if other points of difference are present,
would not preclude the maintenance of each as a separate genus.

Subge7iera. — Two subgenera of Aulosteges have been proposed:
Wyatkina Fredericks.

Strophalosiina Licharew.

These subgenera agree with Aulosteges in essentials, but differ in minor
details.

Wyatkina Fredericks (1931, p. 210 footnote) with subgenoholotype
Aulosteges gigas Netschajew includes : —

“Stropralosia-\\ke forms^ yet with primitive apical apparatus
devoid of delthyrial ridges and euseptoid. Ornamentation consisting
of thin spines. Muscle impressions of the ventral valve and structure
of the dorsal valve as in Strophalosia.” (The Strophalosia of this
paragraph refers to the Russian forms, i.e., Aulosteges.)

Licharew suggests (1937, p. 128) that the delthyrial ridg'es are probably
only x^resent when the deposition of shell substance has been excessive and
these ridges would be the outward manifestation of teeth. This explanation
seems quite plausible. As few Aulosteges possess either teeth or delthyrial
ridges, Fredericks’s subgenus would include nearly all the members of the
genus. Farther observations are necessary before any conclusion can be
accepted with certainty.

Strophalosiina Licharew (1935, p. 369) with subgenoholotvpe Aulosteges
tibeticus Diener is distinguished from other members of Aulosteges by its
geniculate ])ediele valve and peculiar ornamentation. Anterior to the genieu-
latioii the surface of the shell is covered with radial i>li("ations w'hile on the
visceral disc the plications are replaced by irregular concentric wrinkles
and radially arranged pustules.

Tliese subgenera, Wyatkina and Strophalosiina are closely related.
Indeed, from a study of the literature they seem identical, since according
to Hamlet, A. tibeticus = A. medlicottianus, while Renz says that A. gigas “

K. L. Prendergast.

3()

A. medlic\ottianus\ therefore, since those things which are equal to the same
thing are equal to one another: —

A. tibeticus == A. giga-^,
therefore Wyatkina — Strophalosiina.

If the ])remises are true Strophalosiina is a synonym of Wyatkina. The
author has not been able to see the original description and figures of
A. gigas, so cannot discuss its affinities with A. tiheticus. The diagnostic
feature of .^1. tiheticus is the ornamentation, and later writers on A. gigas
do not describe that shell as radially plicate.

Until further information is available, Stroplialosuna and Wyatkina are
accepted as subgenera of Aulosteges.

Aulosteges spinosus Hosking.

PI. iv, figs. 7-10.

1981 . — Aulosteges spi}ws-us Hosking, p. 17, pi. iii., figs. 7a-d,

1938 . — Aulosteges spi7iosus Hosking, p. 37.

Material. — Aus. Mus. F 38444, 37710, Wandagee Station, Minilya River,.
North-West Division. Wandagee Stage.

Diagnosis. — Shell small to medium in size, subquadrangular, wider than
long. Surface of both valves with small perforated tubercles bearing small
spines.

Pedicle valve convex, not swollen, faint median sinus. Umbo high, not
overturned, Area broad, slightly concave and recurved to cardinal margin^
j^seudo-deltidium narrow, highly arched.

Brachial valve flat, triangular extension of cardinal margin large.

Internal features unknown.

Description. — For complete description of this species reference should
be made to Hosking (1981, p. 17 et seq.).

Two rather dissimilar sj)ecies from the North-West Division are here
referred to this species.

Dimensions.

Length

Type :
18-8

38444.
45*1 -f

37710.

20

Breadth ....

19-7 +

43-0 -f

22

Length of hinge-line

14-5

21-3 +

15

Thickness

—

7-4

F 38444 is a com])Iete specimen with both valves. The umbo is distorted
by a cicatrix of attachment and twisted slightly to the left. The area is high
and very worn so that no growth lines or transverse striations are prescr\'ed.
It has the same recurving to the cardinal margin as noted by Hosking in
the description of the holotype. The pseudo-deltidium is narrow and parallel-
sided; it has twisted, with the twisting of the umbo, to the left. An indenta-
tion of the valve, as the beginnings of a median sinus, occurs near the
anterior margin. The ornamentation of the pedicle valve is of fine forward-
projecting spine bases.

Permian Productinae and Strophalosiinae of W.A.

37

The brachial valve is fiat and is without an upturned margin. The
median septum crosses two-thirds of the length of the shell. The surface of
the valve is roughened by weathering, but numerous pits indicate the former
presence of spines.

A much larger specimen, F 37710, is a pedicle valve with a high undis-
torted area. The valve has the same recurved margin and ornamentation as
seen in the type. The median sinus is well-developed anteriorly, but is faint
on the visceral portion of the shell. The area is wnde and longitudinally
striate; the pseudo-deltidium is narrow, parallel-sided and bears nodular
perforated tubercles; it is short, continuing for only half the width of the
area so that a large open space occu])ies the centre of the cardinal margin
(filled, when the brachial valve is in position, by the triangular extension
of the cardinal margin). This specimen differs from the type in size and
in the possession of deltidial spines. Although no intermediate forms are
known it is im])ossible to separate the specimen on those grounds alone.
The deltidial spines are rarely present in species of Aulosteges and their
absence may be due to removal by weathering rather than to a difference in
development.

Bey^iar'ks . — The distinctive feature of this species is the sharp recurving
of the area, a feature not seen in other species of the genus.

The a]^pearanee of a larger shell with the same characteristics as those
described for Aulosteges spinosus nullifies the suggestion of Hosking (1931^
p. 19) that this species was possibly only the young form of Aulosteges ingens
Hosking,

STROPHALOSIA King 1844.

1844. — King, Ayu). Mag. Nat. Hist., XIV, p. 318.

Gniotiipe. — OrihU-' cxcavata Geinitz, Ueber einigc Petrefacte des Zeclisteins
iiiul Miiscdielkalk's. .V. Jh. Min. Geol. Falllont. 1842, p. 578 Taf. x hgs. 12, 13.

Diagnosis. — Small to medium-sized shells. Concave to platy-convex dorso-
ventrally with Avell-developed areas on both valves. Umbo distorted by cica-
trix of attachment. Ornamentation essentially lamellar, lamellae usually
interrupted by spines on pedicle valve; spines rarely present on brachial
valve.

Pedicle valve with teeth litting into sockets of bachial valve. Brachial
valve Avith median septum, non-dendritic muscular impressions and pro-
nounced brachial impressions. Trifid cardinal process well-developed.

Description. — The members of the genus Strophalosia are never large
shells; they vary in shape from longitudinally to transversely oval, with the
hinge-line equal to or slightly less than the greatest Avidth of the shell. The
pedicle valve is usually evenly convex, rarely geniculate. The brachial valve
is, however, usually geniculate, being flat or concave in the visceral portion
AA’ith upturned lateral and antenlor margins. Both valves possess a distinct
area, that of the pedicle \'alve being the larger. There is no ojmn delthyrium,
a pseudo-deltidium being ahvays present.

Strophalosia' has been described as a spinose Productid with an area.
This description is, hoAvever, a little restricted, as all species of Strophalosia^
for example Strophalosia kimberlcgemis mihi, do not possess sjunes. The
characteristic of the ornamentation is rather its lamellar structure. The
lamellae are usually distinct, their outer surfaces and edges forming the ex-
ternal shell surface. This distinctness of the lamellae indicates that in the

38

K. L. Pren'dergast.

members of the genus shell formation was limited, probably by physical
conditions, to certain times of the year, the periods of shell deposition being
separated by intervals of (puescence during which no shell was deposited.
That this response to physical conditions is a generic characteristic rather
than due to abnormal physical conditions may be inferred from a study of
forms living under the same physical conditions in which the lamellar struc-
ture is not seen.

Spines are often present; these may be recumbent or erect, many oi' few;
they may be of two series or uniform in size, and are usually present on
the pedicle valve, though but rarely found on the brachial valve.

Internally, the structures of the brachial valve are characteristic of the
genus, and the changes within the genus are largely reflected in the structure
of this valve. A median septum is always present, varying in length from
one-third to two-thirds of the length of the visceral portion of the valve. It
separates the non-dendritic adductor muscle impressions which vary in posi-
tion but shoAv no relation in their variation to the length of the median
septum. The adductor impressions are usually divided into anterior and pos-
terior adductors, but sometimes (e.g. Bramtonia) no such division is seen.

The brachial impressions arise at the side of the adductor muscle scars
and vary in position with the jmsition of the muscles. Where the muscles
are medial or lower in the valve, that is, towards the anterior margin, the
brachial impressions run almost parallel to the hinge-line and curve abruptly
to the median septum, the condition seen in the Productus group. Where the
adductors are situated higher in the valve the brachial impressions are found
near the cardinal and lateral margins and approximately parallel to them.

Figure 6.

Internal Features of Brachial Valves — to illustrate the alteration in position
of the Brachial Ini])ressions. (a) Productus latirostratus, (b) Strophalosia
lamellosa var. humbletonensis, (c) Str. jukesi, (d) Str. kimberleyensis. (a),
(b) and (c) after Davidson (Xli 2 )-

The cardinal process, while it varies in minor details, retains throughout
the genus, its typical trifid character. The process is always a compact one,
the three parts being on the same horizontal level, in contrast with some
Productids, e.g., Taeniothaeruf^ snbquadratus, where the central lobe is raised
above the lateral lobes. The structures at the base of the process vary;
this variation will be discussed later (p. 30).

In the ]')edicle valve the adductor impressions are non-dendritic, central
and varying in their position in the valve as do those of the brachial valve.
The diductor impressions surround those of the adductor muscles and are
longitudinally striate.

In life the shells of this genus were probably orientated with the heavier
pedicle valve resting on the substratum, the curvature of this valve and the
upturned margins of the bracliial valve keeping the open edges of the shell
above the level of disturbed water. The umbonal cicatrix shows that the

Permian Producttnae and Strophalosiinae of W.A.

39

shell was attached at some period, but it is not known for what length of its
life the shell maintained this position. If the shell was fixed only by cemen-
tation the period of attachment was probably short as no specimens are
known with the shell in this position nor are they usually broken across the
umbonal region as one might expect. If, however, a process of the mantle
aided cementation during life the shell would fall after the death of its in-
habitant. There are no indications to show that such a process or processes
existed, and one must conclude, in the absence of evidence to the contrary,
that the Strophalosias were attached only during the earlier stages of growth.

Remarks. — The large variation within the genus StrophaJosia is un-
doubtedly due to the rate of adaptation in the different species to the seden-
tary mode of life.

In StropJialosia lamellosa var. linmbletonensis the shell shows a clear re-
semblance to Productus in the arrangement of the musculature and brachial
impressions. The adductor muscle impressions are central and the brachial
impressions have a distinctly Productid aspect (see fig. 6, p. 38).

If a form such as this assumed a sedentary mode of life with the point
of attachment beneath the umbo, it is obvious that the structure is mechanic-
ally unsound, as the muscles in -working, will be pulling against the point of
attachment. To overcome this instability the muscles would tend to move
backwards in the shell until they came to rest over the scar of attachment.
This posteriorly directed movement of the muscles would push all those struc-
tures posterior to them backwards, and these would thus tend to atrophy or
be replaced by the originally more anterior elements. The backward move-
ment of the muscles necessarily causes an alteration in the position of the
brachial impressions, as already seen (see p. 38).

In StropJialosia species with anteriorly placed muscle impressions, two
oblique ridges arise, one on each side of the median septum, beneath the umbo
and continue antero-laterally at an angle of about 45° to the hinge-line. As
the muscles move posteriorly these ridges become displaced and gradually
become the horizontal cardinal ridges as seen in Str. Jcimberleyensis mihi.
They also impinge on the hinge-line so that the base of the cardinal process
has different appearances according to the stage of movement of the muscles.

Licharew (193*, p. 12 * ) states that the extreme condition of this move-
ment is that shown by the Russian specimens in which the ridges have been
pushed beyond the hinge-line, taking with them the sockets. The species to
which Licharew refers, Str, fragilis and Str. horrescens, have dentritic ir.uscle
impressions and must be placed as members of the genus Aulosteges. More-
over, examination of the cardinal process of Str. horrescens shows that it has
the typical trilobate structures and is not bifid, as might be expected if its
mode of formation was as stated by Licharew. The backward movement of
the ridges does change the form of the base of the cardinal process, but I
have seen no specimen in which they replace it. The poor condition of the
specimens of Str. clarkei figured by Etheridge (1880) led Licliarew to think
that this series of changes took place within that species. Str. clarkei does
represent an advanced stage, in that the muscle impressions are high and
the ridges have come to assume a iiosition almost parallel to the hinge-line
but the cardinal process has not been affected and the dental sockets are” deep.

Comparison with other genera. — Strophalosia may be distinguished from
both Aidosteges and Productus sensu lato by many features. In general,

40

K. L. PRENDERGAST.

Strophalosia has a well-developed area on each valve, Aulosteges on the
pedicle valve only, while ProducMis is without a true cardinal area. This
distinction does not, of course, hold for all the Produetiis subgenera, in some
of which, a cardinal area is developed. The non-dendritic adductor muscle
impressions serve, however, to distinguish Stroi)}ialoMa from both Aulosteges
and Productus as in tliese genera the adductors are alwaj^s dendritic. The
posterior movement of the muscles and consequent change in pattern of the
brachial impressions is also a typical Strophalosoid feature.

Suhg(*ner a.— In recent years there has been much work done on tlie

species of this genus, and it has now been subdivided into five subgejiera : —

ETHERIDGINA Oehlert (— LEPTALOSIA Dunbar & Condra)

STROPHALOSIINA Licbarew

STROPHALOSIELLA Lii-harew

WYNDHAMIA Booker

HETERALOSIA King

and with Aulosteges as a possible sixth or alternatively as a synonym. The
relation of Aulosteges and Strophalosia has already been discussed (p. 34).

Etheridglna includes those Strophalosias in which the shell is attached
throughout the life of the animal, both by embracing spines and directly by
the pedicle valve. A small cardinal area and teeth are sometimes, though
not always present. Leptalosia, to include those forms possessing an area
and teeth, cemented directly by the pedicle valve, was founded by Dunbar and
Condra in 1932. Examination of the type specimens of Etheridgina com-
plectens (Eth. til.), the genotype of Etheridgiue shows, however, that some
of these specimens possess the hinge structures thought to be typical of
Leptalosia. Leptalosia becomes, therefore, a synonym of Etheridgina. This
question is more fully discussed under Etheridgina.

The genotype of Strophalosiina Licharew is Aulosteges tihetius Diener.
This species undoubtedly belongs to Aulosteges and is, under this system of
classification, regarded as a subgenus of Aulosteges,

Strophalosiella Licharew with genotype Strophulosiella coraeformis
Licharew is a very doubtful genus. The specimen figured and described by
Licharew is a plicate form (“a Strophalosia with Linoproductid ornamenta-
tion^') with well-developed areas on both valves. In the absence of any
knowledge of the internal structure it seems impossible to assign this speci-
men to the genus Strophalosia. Until this knowledge is available it would
seem better to maintain it as an independent genus of unknown affinities.

Wyndhaonia was founded by Booker in 1929 for the reception of a group
of Strophalosoid forms with adductor muscles undift'erentiated into anterior
and posterior elements and without evidence of a period of attachment. The
position of the muscle scars (]daced posteriorly in the valve) would indicate
that the species of this subgenus (TE. vallda Booker and IE. dalwoodensis
Booker) had attached ancestors, and had retained some of the ancestral
characteristics, although they themselves had ceased to be attached. The
assumption of these distinctive features, undifi'erentiated adductor muscle
scars and absence of a cicatrix, is sufficient to characterise the subgenus. The
type species of the subgenus is Wyndhamia dalwoodensis Booker from Per-
mian beds of Branxton, New South Wales.

In the same j)aper (1929, p. 30) Booker also proposed a new subgenus
Bran.rtonia of Productus. The author has not been able to see the specimens
of the genotype, Branxtonia typica Booker, but from Booker^s own figures
and similar specimens from Western Australia, suspects that the specimens

41

Permian Productinae and Stropiialosiinae of W.A.

are Strophalusias. Tliey show the same strueture and arran^'ement of the
musele scars in the brachial valve, and the impression of dental callosities
may be seen on the casts. The s])ecimens do not show an ar(‘a, but until
specimens with some shell preserved are seen it is impossible to say that its
absence is not due to tlnckening' of the hin^'e-line behind the area. If no
area is present then Bi'iui.rionia must be rei>‘arded as an altaclu'd form or
of a line of attached ProcUudids with conse((uent muscle niov(*ment as in
Strophalosia.

IJetf^ralosia Kin<>' (IIKIS, ]). 27H) with genatyi)C IT. slocoitu Kino; is dis-
tinguished from Stropitalosia s.s. by the possession of a non-spinose brachial
valve. It seems doubtful whether a division of the Strophalosias based on the
ornamentation of one or the other \'alve is of great advantage, and although
of the s})ecies here described S. etheridgei and gerardi (?) would be ])laced
in Ileteralosia 1 have left them in Strophalotiia s.l. If King’s division is
accepted, a new genus, without spines on either valve, must be proposed to
include S. clarkei and kimherleyensis.

Strophalosia cf. Strophalosia beecheri K’owley.

IM. Vj tigs, l-.'i.

isn.‘h — Sh'o; lialo.s'ia brrchfvl Kowley, ]). 308, ])1. 14, tigs. 18-19.

\\H)S~~Stro/)haliosi(i hcrclieri Kowley, ]>. 70, pi. 17, tigs. 24-25.

1914. — Sirophalosia hccchcri KoAvley, Weller, ii. 140, pi. xix, tigs. 37-38-

Material. — U.W.A. 20449, one s})ecimeU" -conjoined valve; Waltliarrie Pools,
Wooramel River District; Callytliarra Liinestoii(‘.

Ddtgnosiy . — Shell small, almost circular in outline, hinge-line slightly less
than greatest width of shell. Surface oiiianumtation of ei'(!ct s[)ines of
approximate (luim-umual arrangement and tine growth lin(‘s.

Pedi(*Ie vahe convex, asymnudric without sinus though flaitened medially
and sloping stt'eply anteriorly aiul latei'ally. Cicatrix large, placed on one
side of umbo. Area as long as hinge-line, Hat and trianguhu’. Pseudo-
deltidium triangular. Ttadh short and diverging.

Brachial valve concav(g area distinct, almost as large as that of [)edi(de
valve, pseudo-cliilidium larg(' and triangular.

internal features not known.

Descriptio}! : Dimensions of Bpeeimen,

Height l*2.r>

Maximum breadth .. .. 12.(i

Length of hinge-line . . . . 8.8

Specimen too crushed to give accurate measurcmient of thickness.

The pedicle \-alv(‘ of this specimen is asymnudric du(‘ to the largo cicatrix
of attachment on on(> side of the umbo. It is evenly swollen in tin* middle
wiih the sides and the front sloping steeply giving it a somewiiat s(iuashed
appearance. The ears are small and tint with (d)tuse cnrdinal angles. The
specimen is iinl'ortunately badly weathered so that the ornamentation is
ol)scured. Slender erect si)ines with ai)proximate (piincuncial arrangement
are the obvious feature of the ornamentation, rhe concentric shell laminae
<d‘ the original slu'll surface, if these Avere prt'sent, being ()bs('ured. The

42

K. L. Prendergast.

area is flat and lies in the plane of the commissures; it is interrupted by a
triangular slightly convex pseudo-deltidium. Owing to the crushing of the
brachial valve, rhe teeth are seen to be short, stout and diverging.

Even in its uncrushed form, the brachial valve must have been very
concave. The ornamentation cannot be seen at all unless some pits due to
■weathering indicate that it was spinose. The area of this valve is as large
as the pedicle area, with a well-develoi>ed pseudo-chilidium.

Comparison n'itJf other species. — This specimen resembles Strophalosia
heecheri Rowley (Weller 1914, p. 146, pi. xix, flgs. 37-38). Weller, in his
description of this species, quotes its asymmetric character as a diagnostic
feature. From his flgure 37 it Avould appear that in that species, as in this
specimen, the asymmetry is probably due to the cicatrix on one side. It is,
however, a broader form than the present species and Weller says broader
than long: but as he, too, had only one specimen, the variation within the
species is not known.

Bemarhs. — This specimen has been compared to Strophalosia heecheri^
but not identifled finally as a member of that species since further specimens
are required, both from America and Western Australia, before any definite
identification is possible. More specimens, too, may establish the present
specimen as a member of one of the variable Western Australian species.

Strophalosia clarkei (Etheridge).

PL V, fig. 4.

l^'i'I.—Productus clarl-ei Etheridge, p. 334, pi. xvii, figs, 2, 2a, 2b; pi. xviii
figs. 4, 4a. 1 , ^ , i ,

1877. — ProduePns clarl-ci Etheridge, de Koninck, p. 203, pi. x fio-. 5 * -.q xi

fig. 3. X , 1 , ^ , i . ,

1878. — ProducUis eJarhei (Etheridge), Etheridge, jun., p. 51.

1880. — Strophalosia rJarl'ci (Etheridge), Etheridge, inn., p. 289, pi. ix, figs.
18-21; pi. X, figs 22-28; pi. xi, figs. 29-31; pi. xii, figs. 32, 33.

1892. — Strophalosia clarhei (Etheridge), Etheridge & Jack, p. 258 pi. xiii,
figs. 12-17; pi. XIV, fig. 19. ’

Iloloti/pe, helonged to Daiiitree Collection. Present, location of type specimen
not known.

Other Material. — B.Al. B 5885, Capertree, N.S.W. (figured by Etheridge
1880). Ans. Mus. F 36234, 36235, .liniba Jiniba. near Mooka Springs, North-
West Pivision. Gascoyne series.

Diagnosis. — Shell transversely oval to subquadrate, hinge-line less than
greatest shell width. Pedicle valve inflated, without median sinus. Brachial
valve fiat to concave. Ornamentation of silky laminae on both valves.

Interior of pedicle valve with thin median ridge sti.arating dendritic
adductor muscle impressions. Diductors large, fiabellate and longitudinally
striate. Teeth large.

Brachial valve with strong cardinal process continued into long median
septum. Dental sockets large and oblique. Adductors high in valve. Brachial
impressions .strong, folloAving edge of valve; valve geniculate with marginal
rim perpendicular to visceral disc. Line of three depressions (adductor scars,
depression below muscle scars and ends of brachial impressions) on each
side of median se]>tum.

Permian Productinae and Stropiialosiixae of W.A.

43

Description. — Little can be added to the descriptions of this species in
Etheridge (1880) and Etheridge and Jack (1892). The two specimens from
Jimba Jimba are brachial valves whose external features are obscured by
matrix. The internal features are as described by Etheridge. The charac-
teristic structures of the valve are the line of three depressions (those of
the adductor impressions, depression below the muscles and ends of brachial
impressions) on each side of the median septum and the groove and marginal
ridge on the outside of the brachial impressions. Inside the brachial
impressions the shell is raised and prominent.

C omparison with other specjes. — The differences between this species and
Strophalosia himberleyensis mihi will be found under the heading of the
latter species.

Etheridge (1880, jn 290) suggested that Strophalosia gerardi King and
Strophalosia clarhei (Etheridge) were possibly one and the same species.
The absence of any I’eliably identified specimens of Str. gerardi showing the
internal characters led him to postpone a final decision in the matter. The
situation of Str. gerardi is still the same, but these species may be distin-
guished by their external characteristics. Strophalosia gerardi has a relatively
much shorter hinge-line, and its external ornamentation of obli(iue adherent
s])ines on coarse concentric lamellae distinguishes it from the smoothly silky
surface of Str. clarhei. Etheridge (1872) described the ornamentation of
Str. clarhei as “covered with fine, vertical, wavy lines, projecting from
which are numerous slender spines.” This ornamentation 1 have not seen,
weathering having removed apparently both the vertical lines and the spines.
However, the descrijition would certainly not fit the coarse adherent spines
of Str. gerardi.

Strophalosia etheridgei n. sp.

PI. V., figs. 5-12.

Sifnti/pcs, — O.S.W.A. 1/.T242 (a), creek, mile Avest of Callytharra Springs,
Wooramd Piver. Callytliarra Limestone.

Topol ypos. — G.S.’W.A. 1/5242 (b), same locality and horizon.

OIhrr Mafcrial~V.\Y.A., 20267,’ Possil Cliff, Tnvin River. Pos.sil Cliff hori-
zon. TT.W.A., 20247, Fossil Ridge, TrAvin River Pistrict. Fossil Cliff horizon.
Alls. Allis. F 3SI6.3. 3S464, 38448, Wandagee Station, Alinilya River, North-West
DiA’isioii. Wandagee stage.

Diagnosis . — Shell small, thin, transversely-oval to almost circular in out-
line. Hinge-line less than greatest Avidth.

Pedicle valve flattish, evenly convex, non-sinuate, area relatively Avide
wilh nari’OAv but not prominent ])seudo-deltidinm. Ornamentation of adher-
ent spines interrupting .silky laminated surface structure. Umbo not promi-
nent, area of aftachment small. Teeth small and diverging. ALiscle scars
deltoid in outline, non-dendritic, almost under umbo.

Brachial valve flat in visceral region Avitb upturned margins. Ornamen-
lation of silky laminae and concentric Avrinkles. When Aveathered, valve pitted
anteriorly. Cardinal iirocess inclined, continued into short median septum.
Sockets prominent, inclined at high angle. Afuscle inpwessions rounded, at
base of cardinal process.

44

K. L. Prexdergast.

Description . — The small size of this species is shown by the table of
dimensions : —

—

Maximum

width.

Length

hinge-line.

Height.

Synlypes :

1. Conjoined valves

8*0

4-6

6-5

2. Pedicle valve

8*5

5* 7

7-0

3. Brachial valve ....

6-7

5-0

5*4

4. Conjoined valves

9-0

0-2

8-0

Irwin River Specimeyis :

1. Fossil Ridge

10-8

6-8

8-5^

7*1 ^

4-2

7-1 J

Brachial

2. Fossil Cliff

10-4 4

5-0

9-4 j

Valves

8-9 +

5-3

8-4

15-9

11-0

15-4

Pedicle

\"alve

The i)ediele valve is small, thin-shelled and moderately inflated; it
diminishes in convexity gradually towards the lateral and anterior margins,
somewhat suddenly towards the cardinal margin. The area is small, narrow
and triangular, its lengtli less than that of the hinge-line. The shell sub-
stance is lamellar and has a silky texture, the laminae interrupted by the
spines, which closely adhere to the surface of the shell. A few erect spines
are present on the ears. The muscle scars are situated high up in the valve
and have a deltoid outline, the delta divided medially l)y a groove into the
two triangular areas. They are non-dendritic.

Cardinal Process of Stroplialosia etlieridgei in sp. (XH).

The brachial valve is flat in the visceral region with geniculated margin
and has a silky laminated and imbricated surface. The hinge-line is short
and the cardinal angles are obtusely rounded. The area is distinct, triangular
with the umbonal angle flattened; the pseudo-chilidium is triangular and
wide with two deep, narrow sockets, inclined to the median septum at its
base. The cardinal process is narrow near the hinge-line, but swells out to a

Permian Productinae and Strophalosiinae of W.A.

45

bulbous top. This is, however, usually worn down so that the process appears
to continue posteriorly at an even width. The semi-circular visceral portion
■of the brachial valve is interrupted by the short median septum, which is
.swollen between the rounded muscle impressions. The valve is hollowed out
l)elow the sockets and in a direction parallel to the hinge-line, a ridge sei>arat-
ing this hollow from the cardinal margin.

Compariso7i with other species. — The small size of these specimens dis-
tinguishes them from any species of Strophalosia so far described from West-
ern Australia. They were labelled by Miss Hosking in the Geological Survey
■Collection, as being young specimens of a new species. The geniculated
margin of the brachial valve indicates, however, that they are probably in
the adult or nearing the adult condition unless they are the young forms of a
Prohoscidella-V[kQ Strophalosia. In the absence of any shells which might
be assigned to such subgenus I have described them as adult forms.

Many features of these shells are also common to Strophalosia clarkei
Eth. til. and Strophalosia gerardi King, although they cannot be regarded
as closely related to either of those species. The brachial valve particularly,
with its silky lamellar structure, short hinge-line and rounded alar angles,
resembles S. clarkei. The pedicle valve, however, would distinguish them at
once from that species in its ornamentation showing a marked resemblance
to S. gerardi; from young members of this species they would be distinguished
by the shape and convexity of the pedicle valve.

Strophalosia gerardi King.

PI. V, fig?. 13, 16, 17.

1846. — Strophalosia grrardi King, p. 93 footnote.

, 1847. — Strophalosia gerardi King, de Koninck, p. 137.

ISoO. — Strophalosia gerardi King, p. 96, pi. xix, figs. 6-7.

1857. — Strophalosia gerardi King, Davidson, Introduction, pi. viii, fig. 211.

1880. — Sirophaloeiia gerardi King, Etheridge, jun., p. 294, pi. xii, figs. 34-37;
■pi. xiii, fig. 38.

1884. — Strophalosia gerardi King, Etheridge, jun., p. 87.

1892. — Strophalosia gerardi King, Etheridge & -Tack, p. 260, pi. xiii, fig. 18;
pi. xiv, fig. 18; pi. xl, figs. 7-8.

1932 . — StrophalosUi gerardi King, Eeed, p. 20, pi. i, fig. 7.

Maferial.^ — Rolofype. Univ. Coll. Galway, I.F.8., Xo. E.C.D. 267.

Diagnosis. — Shell outline transversely oval. Pedicle valve convex, non-
sinuate; brachial valve concave following outline of pedicle valve. Ornamen-
tation of concentric lamellae and adherent tapering sjjines. Area on both
valves with pseudo-deltidiiim and pseudo-chilidium. Length of hinge-line
less than maximum width of shell.

Pedicle valve swollen, with greatest convexity in visceral region, non-
geniculate. Cardinal angles obtuse, ears small.

Bi'achial valve concave, greatest concavity near umbo, shell sloping
thence gradually to all margins.

Internal features unknown.

Description of specimen. — Univ. Coll. Galway, E.C.D. Xo. 267. Viewed
from the brachial side the shell is a regular oval in outline, the transverse axis
passing postero-anteriorly. From the pedicle side it has rather a triangular
aspect.

K. L. Prexdergast.

4()

The pedicle valve is flatly convex, the maximum convexity behind the
umbo, thence flattening gradually to the anterior and lateral margins and
steeply to the cardinal margin. The area is short, equal in length to half the
maximum width of the shell ; it is relatively high with a very narrow, ahnost
straight-sided pseudo-deltidium. The area of attachment is large, and has
the appearance of callus over the shell surface. The ornamentation of the
pedicle valve is essentially spinose, the spines adherent, Avith an irregular
quincuncial arrangement. The structure of the shell surface is lamellar though
the lamellae are largely obscured.

The brachial valve is concave. It slopes steeply from the umbo to the
middle of the shell, thence folloAving the curve of the pedicle valve. The shell
structure is lamellar, but the valve is Aveathered, and no spine bases are seen,
the ornamentation consisting of the truncated lamellae and irregularly spaced
nodules. The area of the brachial vah’e is narroAV but distinct. Internal
features not shoAvn.

Dimensions.

Height .. .. 30.4

Maximum breadth . . . . 38

Thickness .. .. 10.8

Etheridge says (1880, p. 296) that he finds “what appears to be the bases
of insertion of spines’’ on the brachial A’al\'e of King’s type specimen. It
is difficult to come to a decision on this point from a study of the type speci-
men alone as, although there are holloAA^s and small prominences on the
lamellae, it seems almost impossible to decide that these prominences are or
Avere spine bases. They shoAV no concentric structure and no sign of aperture.
It is noticeable, hoAA'ever, that Avhere the spine bases are Avorn down on the
l)edicle Auilve, they, too, show no structure. On his figures of the Queensland
specimens, too, Etheridge figures no specimen Avith spines on the brachial
valve.

Strophalosia sp. cf. Str. gerardi King.

PI. V, fig. 14, 15.

Maicrial. — F.W.A., 12399. One pedicle valve. Fossil Cliff, Irwin RKer.
Fossil Cliff horizon.

Description . — The shell is convex, non-geniculate, Avith the greatest con-
vexity in the visceral region l)ehind the cardinal margin. From this area of
maximum convexity the surface slopes gradually to the anterior and pos-
terior margins, but more steeply laterally. A large cicatrix covers the
uml)onal region. The ai'ea is concave Avith a Avell-marked pseudo-deltidium,
at the base of Avhich arise strong diverging teeth. The length of the area is
about half the maximum l)readth of the shell.

Dimensions.

Maximum Avidth

31.7

Length of hinge-line

. 15.3

Height

27.0

Length of pedicle A-alve

. 38.9

The shell surface is lamellar Avith strong olfiique spines arranged in
irregular concentric roAA’S. Posteriorly ami on the cardinal slopes the spines
are small and adherent; they become larger toAvards the anterior margin.

Permian Productinae and Strophalosiinae of W.A.

47

Internally the adductor impressions are seen high up in the valve; they
<^l•e separated medially by a narrow groove. Longitudinal striations mark
the shell at the side of the adductor impressions and are probably the im-
pressions of the diductor muscles.

Comparison ivith other species. — This specimen has been referred to
Strophalosia gerardi rather than identified with it. The general outline of
the Irwin River shell is much broader than any specimen of Sir. gerardi, and
the spines are more widely spaced and less oblique than in that species. This
.specimen is possibly a local variant of Str. gerardi^ but in the absence of
intermediate forms cannot be satisfactorily united with it.

Strophalosia jukesi Eth. fil.

P1..V, figs. 18-20.

f8S0. — Strophalosia jukesi Etli. fil., p. 307, pi. xiii, figs. 39-43.

1888. — Strophalosia jukesi Eth. fil., Jolmstoii, pi. xiv, fig, 7.

Si/ntypes. — R.M. 96874, 96931. New South Wales. Exact locality and hori-
zon unknown. Figured by Etheridge 1880, pi. xiii, figs 39-43.

Other Material. — Commonwealth Palaeontologists^ Coll. P. 14. Talbot’s
<7011-11, Woorainel River District. U.W.A., 20264. same locality and horizon.
U.W.A., 20266, same locality and liorizon.

Diagnosis. — Shell elongate-oval in outline, hinge-line less than maximum
shell width. Ornamentation of spines and concentric lamellae on both valves.

Pedicle valve evenly convex with flattening towards anterior margin
and steo]) slope to cardinal margin. Area wide and distinct, pseudo-deltidium
triangular. Ears small. Cardinal angles obtuse. Muscles high in valve,
adductors central and compact, diductors tlabeliate and longitudinally striate.

Brachial valve concave, greatest concavity in front of umbo. Area
distinct, half as wide as area of ]:)edicle valve. Adductor muscle impressions
divided into anterior and posterior sections. Cardinal process large, trifid,
continuing into strong median septum.

Description. — A number of ferruginous casts have been referred to this
species, as they agree in all particulars with Tasmanian specimens. For a
<-:omplete descri])tion of the species see Prendergast (Proc, Rog. Soc. Tas-
mania).*

Strophalosia kimberleyensis n. sp.

PI. vi, figs. 1-5.

1890. — Strophalosia cJarkei Etheridge, Foord, p. 103, pi. v, fig. 7-8, text-fig. 6.

1903. — Strophalosia sp. ind, Etheridge, .inu., p. 20, pi. i. figs. 10-12.

llolotype. — F.W.A. 20452 north of Hill C, south side of Grant Range, Kim-
berley Division. F])])cr Ferruginous Series.

Paratypes. — T'.W.A. 20460, same localitv and horizon.

ir.W.A. 2045.5, Nooncanbah Homestead, Kimberley Division. Nooneanbah
Series.

Topotype.s.—V.\\\A. R 127.

Other Material.— P>.M. B 4590, 4591, south-east of Mt. Abbott, on Fitzvoy
River, Kimberley Division. Horizon unknown. (Figured by Foord 1890). W.A.
Mus. F 166 (47441, Cookilya Pool, Wandagee Station. Minilva River. Wandagee
stage. (Figured by Etheridge, 1903). W.A. Mus. a 12, south-east of Mt.
Abbott, Fitzroy River, Kimberley Division. Horizon unknown. Aus. AIus. F
37523-37529; IMinilya River. Wandagee stage. Aus. Mus. F 36221-36225, Jimba
Jiml)a Station, Gaseoyiie River. Byro stage. U.W.A. 13. 51. Nooncanbah Home-
stead, Kimberley Division. Nooncanbah series.

Diagnosis. — Shell sub(]uadrate to ovate in outline, pedicle valve inflated
to bemisi)herieal. braehial valve evenly concave, liinge line almost greatest
width of shell.

48

K. L. Prexdergast.

Pedicle valve with lamellar concentric ornamentation ; cardinal area
elongate triangular with small triangular pseudo-deltidium. Internally Avith
strong teeth, valve thickened postero-laterally with strong ridges bounding
visceral portion laterally. Adductor muscles non-dendritic, narrow, on each
side of median line; diductors longitudinally striate arising in front of
adductors and extending from near middle line to lateral margin. Muscles
set in distinct fossae.

Brachial valve with ornamentation as in pedicle valve. Valve thickened
anteriorly, regularly concave exteriorly but geniculate interiorly. Cardinal
process trifid, strong, Avith sockets ax base; sockets separated by diagonal
ridges from depressed muscular area. Posterior adductors elongate laterally,
anterior adductors elongate antero-j^osteriorly. Brachial impressions arising
from lateral edges of posterior adductors, following lateral and antero-lateral
margins then turning backAvards toAvards median septum. Septum continuing
about two-thirds length of valve. Cincture separating visceral disc from
trail.

Description of Holctifpe. — The holotype is an almost complete specimen
Avith both A’alves in position. The pedicle A^alve is unfortunately weathered so
that the external shell layers are absent. The valve is thick and of lamellar
structure. The area is long and triangular, its Avidth equal to one-fifth of its
length; it has a narroAAq closed, triangular delthyrium. The brachial valve
is regularly concave and is ornamented by concentric lamellae, the edges of
the lamellae forming the shell surface. It has a narroAv area disposed almost
at right angles to that of the pedicle valAT.

Description of Daratypes . — The paratypes are Iaa^o specimens, one a
pedicle valve, the other a brachial valve, selected to shoAv the internal
features of the species.

In the pedicle valve (20400) the teeth are strong and du'erging from
the hinge-line; laterally they are joined to a projecting ridge separating the
ears from the visceral caA’ity. The shell is thickened in the region of the
ears, the ridge continuing as a platform to the lateral margin, its continuity
being broken up by a narroAv groove behind the ridge. This groove serves
as a socket, a ridge on the brachial valve fitting into it. The muscle marks
are distinct, the adductors elongate on each side of the middle line, arising
almost under the teeth. The diductors are large, longitudinally striate, and
anterior to the adductors they are separated by a rounded prominence in
the middle line.

Figure 8.

Cardinal Process of Strophalosia kimberleyensis n. sp. (X3).

Permian Productinae and Strophalosiinae of W.A.

49

The interior of the brachial valve (20455) is geniculate, due to thickening
of the lamellae in this region, a narrow groove (cincture) separating the
visceral disc from the trail. In the visceral part the valve is almost divided
by the median septum which continues forward for two-thirds of the length
of the visceral disc; it is continued posteriorly as a trilobate cardinal process,
at the base of which are deep sockets. The adductor muscles are below the
sockets and set in a depression which gradually decreases anteriorly (this
depression is bounded posteriorly by a ridge parallel to the hinge-line). The
anterior adductors are elongate postero-anteriorly. The posterior adductors
are elongate laterally and from their lateral extremities arise the brachial
impressions. These continue the line of the ridge below the sockets until
they almost reach the lateral border, they then follow the lateral and antero-
lateral margins of the visceral disc and turn upwards parallel to the median
septum. They cannot be traced in this specimen beyond the level of the
end of the median septum.

Dimensions of Shells.

Height.

Curvi-
linear
length of
pedicle
valve.

Length
of hinge-
line.

Max.

width.

Thick-

ness.

Holotype

28-2

56-0

22-0

27*8

10*8

Paratypes —
20460

19-2 +

31-0-h

23-7

30*0

20455

24-3

24-5

29*3

Brachial Valves

29-3

29-4

35*7

25-0

25*24-

28*1

Combined Valves

24-9

43-0

23*9

28*5

14*5

29-0

46-0

25*9

28*8

14*4

25*0

42*0

24-5

28*7

9*8

As may be seen from these figures, the height and width of the valve
are the same in full-grown specimens, but as the maximum width is situated
in the posterior half of the shell, younger specimens are wider than long.

The pedicle valve varies in shape from quadrangular to almost semi-
circular and in convexity from weakly convex to hemispherical. As the shell
becomes more strongly curved with age, the convexity is an indication of
the age. There is, too, a difference in shape of some of the brachial valves
due to the varying amount of thickening anteriorly; thus the value may be
regularly concave or slightly geniculate.

Owing to decortication of the shell surface, it is impossible to describe
the tiue ornamentation of the shells. The lamellae are distinct on both valves
and some specimens of the brachial valve show that these are crossea Dy
very fine radial striae. A iew only of these are seen, but they are continuous
from one lamella to the next. One pedicle valve has, also, a small patch of
radial ornamentation preserved. It is ]>ossible, therefore, that, were the shell
surface preserved, the shell could be described as radially striate; alterna-

;50

K. L. Prendergast.

tively the radial markings may be the result of weathering. No pits^ such
as would indicate the presence of spines, have been observed and it would
seem, therefore, that this species is non-spinous.

The variation within the species is also reflected in the internal structure
of the brachial valve. The median septum may be equal in length to half
or increase up to two-thirds of the length of the visceral disc. The brachial
impressions may follow closely the lateral margin or be placed some distance
from it.

Bemar'ks . — The most peculiar characteristic of this species is the nature
of the visceral cavity, the thickened region of the ears and the ridge projecting
into the cavity. This may be seen externally, when the brachial valve becomes
squashed in, showing the shape of the cavity.

Comparison n'ith other species. — I regret that I have been unable to
trace the type of S. clarkei (Eth.), which belonged to the Daintree Collection.
Externally this species resembles S. clarkei^ though it differs in many respects,
as Etheridge (1903, p. 21) has already pointed out. With more and better
specimens many of these distinctions cannot now be maintained, but this
species, S. kimherleifensis is distinct in the depressed umbonal region, rela-
tively longer hinge-line and concave rather than flat brachial valve. Internally,
it occupies a ])osition intermediate between S. clarkei and S. jukesi. In
S. clarkei the median septum is as long as the visceral disc and the bracliial
impressions are transversely elongated; in S. jukesi the septum is short and
the brachial iinjiressions narrow. Thus in S. kimherlei/ensis with short median
septum and wide brachial impressions we see a species with characters of
both the other species. Externally the species is distinct from S. jukesi and
-8. gerardi in the absence of spines.

Strophalosia multispinifera n. sp.

PI. vi, figs. 6-8,

Holofype. — U.W.A. 204.58. Scarp two miles east of Cliristmas Creek Home-
stead, Kimberley Pivision. Nooneanhah Series.

Topoiypes. — P.W.A, B 74. Same locality and horizon.

Other Material. — I^.W.A. 20456, Hill 0., south of road, Grant Pange, Kim-
berley Division. Nooneanhah Series.

Biagnosis . — Pedicle valve large, swollen, transversely oval in outline,
hinge-line less than greatest width of shell.

Ornamentation of fine oblique spines with regular quincuncial arrange-
ment; group of spines on each ear. Valve Avith elongate area, triangular
delthyrium. Cicatrix small. Teeth large. Brachial valve unknown.

Description. — The pedicle Amlve is evenly convex Avith a slight median
depression, not sufliciemly marked to be termed a sinus. It is 45 mm. high,
Avith maximum Avidth of 49.6 mm., AA'hile the hinge-line is 34.8 mm. long.
The surface of the vah’e is marked by closely spaced sjnne bases arranged
in irreg’ular quincunx; it is ]Arobable that these ga\’e rise to small forward
projecting spines, but no si>ines are preserved on the body of the shell.
The spines have a density of 6 in 10 mm. measured transversely at a distance
of 20 mm. from the umbo. The spines become more closely packed on the
ears to give a compact group at each end of the hinge-line. They are closer
together and more h’regular anteriorly. The area is short, its Avidth approxi-
mately half its length; it is longitudinally striated and interrupted by a

Permian Productinae and Steophalosiinae of W.A.

51

pseiido-deltidium which is not raised above the general surface of the area.
Only one tooih has been seen ; this is large and rounded with the concavity
towards the lateral border.

The umbo is not prominent, having been absorbed by the cicatrix of
attachment; this is small and usually almost perpendicular to the surface of
the area.

The brachial valve is unknowm.

Reynarks. — Within this species there is a large variation in shape, many
of ihe shells becoming elongated. It is noticeable that in this species, as in
many Strophalosias (e.g., S. kunherleifetisis mihi) the shells appear very
narrow’ in the hinge region when seen as an internal cast. This is due to the
very close fitting ears which leave almost no space between those of the two
valves when these are in contact.

Comparisoyi with other species. — This species is characterised by the
regularity of the spines over the body of the shells, the spine groups on the
ears, the small regular point of attachment and the absence of lamellar
structure. It approaches nearest to 8. tenuispina Waagen from w’hich it
may be distinguished by the much coarser spinosity of that species and the
larger size of S. ynultispinifera.

STROPHALOSIA sp. A.

PI. vi, fig. 10-12.

Material, — I'.W.A., 20448 Fossil Cliff, Iiwin Kiver District. Fossil Cliff
horizon.

Diagnosis: — Shell longitudinally oval in outline, evenly curved both
transversely and longitudinally. Hinge-line less than greatest wndth of shell.
Ornamentation of lamellae wnth fine adherent spines concentrically arranged.
Pedicle valve sw’ollen, non-geniculate, sides sloping steeply. Area narrow,
triangular. Small divergent teeth.

Brachial valve unknowm.

Description. — A single jmdiele valve from the Iiwvin River District show’s
distinctive characters.

Dimensions.

Height

11.9

Maximum width . .

11.6

Length of hinge-line

6.8

Length of jTedicle valve

18.6

The valve is swmllen w’ith the hinge-line less than the greatest w’idtli of the
shell : the shell is w’idest near the anterior margin. The umbo is completely
obliterated by a large scar of attachment, it has become incurved so that
the area now’ faces antero-dorsallv. The spines are regularly arranged in
concemtric row’s and are all of equal size, there being no sign of a secondary
series of siunes as occurs in so many of the Indian forms. The rows are
widely separated and between them are the silky laminations of the shell
structure.

The area is well-develoned w’ith a wdde delthvrium: the teeth a*-^ lar^re
and diverging. The muscle impressions are high in the valve. The adductors
are central, separated bv a median ridge and enclosed on the outside by the
diductors, which are flabellate and ]moduced anteriorly.

5:2

K. L. PRENDERGAST.

The distinctive characters are the widely spaced concentric rows of
spines combined with ihe elongated shape of the shell.

Comparison xvith other species . — This shell appears most similar to
■Strophalosia rarispina Waagen (1884, p. 645) but the pedicle valve of that
:species has a slight median sinus and the spines are in two series.

Strophalosia tenuispina Waagen.

PI. vi, fig. 9.

1884 . — Strophalosia tcjuiispiiia IVaagen, p. 654, pi. Ixiv, figs. 2-7.

Material.— V.'W. A. 20451. One pedicle valve. Fossil Cliff, Irwin River.
Fossil Cliff horizon.

Diagnosis . — Shell circular to oval in outline, hinge-line equal to maxi-
mum width, pedicle valve but slightly inflated, dorsal valve concave. Surface
ornamentation finely lamellose with sparsely distributed oblique spines.
Pedicle valve non-sinuate, area high, with Hat, narrow linear pseudo-delti-
dium. S 2 uall scai' of attachment.

Brachial valve with small linear area, surface ornamentation not known.

Descrigytion . — This species is known by a single specimen of a pedicle
valve fi'om the Irwin River.

Dimensions of Valve.

Height . . . . . . 17.81

Afaximum width . . . . 21.81

The specimen is rather weathered, but shows the fine spines; these are
represented now only as rather Avorn bases; the spines are all tangential to
the shell surface and some of them adpressed. They are arranged more or
less ill concentric roivs, each row separated from the previous one and that
folloAving by lamellae. On the lateral margins larger spines are develojied,
and these show a curious marking, almost a facet developed on the ventral
surface of each spine. Whether this is the effect of weathering it is im-
possible to say. The spines, apart from their concentric arrangement, also
shoAv a roughly (luincuncial arrangement.

The area of this species is vei’v distinctive among Strophalosias, being
flat and in the jilane of the commissure of the valves. It is interrupted by a
very narroAV, parallel-sided pseudo-deltidium.

Cowgyarison with other species. — For comparison of this species ivith
Strophalosia etheridgei mihi, see account of that species.

ETHERIDGINA Oehlert.

1887. — Oehlert, in Fischer’s Manuel de Couchylogie, p. 1278.

Genotype. — rrodncfiis complectcns Fth. fil., 1876, On an adherent Form of
Produdf-'us and a small Spiriforina from Scotland. Quart. J. geol. Soc. Lord.,
p. 454, pis. xxiv, XXV.

Diagnosis. — Shell small, concavo-convex, dorso-ventrally, Avith semi-oval
marginal contour; generally as broad as long; hinge-line less than greatest
Avidth of shell. Shell attached to foreign bodies by embracing spines or by
cementation of ])edicle A'ah'e. Surface ornamentation of concentric undulat-
ing Avrinkles. Area primarily present, Avith dental sockets and teeth. Muscle
scars non-dendritie.

Permian Productinae and Strophalosiinae of W.A.

53

Description. — This genus was tirst described from the Carboniferous of
.Scotland and is now also known from the American Carboniferous and the
Permian of Australia. The shells are always small and vary in their mode
of attachment with the nature of the foreign body which acts as host. Thus,
in the Scottish Carboniferous forms, where they occur more usually on
crinoids, the spines along the hinge-line are greatly elongated and embrace
the stem which may grow over and completely enclose the brachiopod. In
the American forms (and in the Australian) on the other hand, which attach
themselves to a flat surface, the shell is cemented directly by the whole of the
pedicle valve — the spines along the hinge-line being produced parallel to the
.surface and adhering thereto.

The area, in those forms in which it has been observed, is narrow and
extends along the total length of the hinge-line. In the pedicle valve teeth
may be present or as is more usual in the Scottish forms they are represented
only by vestigial or rudimentary small lumps.

Discussion. — In his original descri 2 :>tion of this genus Oehlert quotes the
date as 1877 ; this was probably an error for 1887. In founding the genus,
Oehlert referred it to ProductuSj it having, he thought, in common with that
genus, the absence of area and of teeth and the j^edicle valve not distorted
by cementation.

The definition of the sub-family StrophaJosiinae by Schuchert (1913)
as productids anchored by spines or by most part of the iicdicle valve,
allowed Etheridgina, like Chonopectiis, to be included in the subfamily with-
out inferring any relationship to Stroplialosia or indeed, any similarity other
than that of habitat. Greger (1920, p. 535) referred a number of small
adnate American species, formerly included in Stropiudosia, to this sub-genus
hut acce])ted it as a sub-genus of Prodnc-tus. Careful examination of the
type specimens of the genotype of this sub-genus shows, however, that ii
should be referred to Stroplialosia rather than to Productus. Several of the
co-types (Royal Scottish Museum, Nos. 5074-5083) show a small though rela-
tively well-develo])ed area and teeth have also been seen on one specimen.
Also the adductor scars of the brachial valve are non-dendritic. These
features clearly distinguish it from Productus and show its relationshijt to
Stroplialosia-. Etheridgina is, therefore, ])laced here as a sub-genus of
Stroplialosia.

In 1932 another productid sub-genus — Leptalosia — was ])ro|)osed by
Dunbar and Condra (1932, 2 >p. 189, 190, 2(i0) to include adnate rroduclids
attach(‘d by the wliole of the ])cdicle valve. The members of this sub-genus
had an area and hinge teeth in the ]K'dicle valve of the older S 2 )ecies (the
sub-genus ranged from Ujiper Devonian to CarI)oniferous) and a distinctive
small jn'oductelliform cardinal ])rocess. The genoty]m is Etheridgina
scintilla (Beecher) ((]uot(*d by them as Stroplialosia scintilla although this
Species had been referred to Etheridgina by Greger in 1920). The sub-genus,
according to its authors, differs from Etheridgina in ijossessing a cardinal
area and in the foian of its cardinal ])rocess the resemblance of the two
genera bdng liomoeomorphic and the result of a similar habit of growth.

It is worth considering these diffei-ences further. As has been already
mentioned, the type sy^ecies of Etheridgina does ptossess an area. Dunl)ar
and Condra (o]>. cit., o. 2fi0) describe the 2 >roduetellifoj’m cardinal prnce-:^
:as ^‘very small and bifid, composed of two narrow posteriorly facing and

54

K. L. Prexdergast.

closely adpressed muscular apophyses supported in front by a pair of very
short diverging ridges separated by a depression.” This clear definition is.
somewhat marred, hoAvever, by their statement (p. 192) that “as now con-
ceived the genus {Productella) is undoubtedly a “Dump-box” for primitive
Productoids and includes the ancestral radicles of several later genera.'^
Which of these radicles, then, have they selected to typify the genus*? In
view of Kozlowski’s Avork (1914) on the cardinal processes of Froduetus
species this is a serious omission, as he has shown quite conclusively that not
only does the cardinal process vary from species to species within a genus,
but also Avith age among the members of any one species. On the other hand,.
Dunbar and Condra may consider that all the species of Froductella,

Avitli its present “dump box” composition, have a primitive type of cardinal
process AA’hich therefore may be taken as characterising the genus; in other
Avords, that Devonian Productoids of AAfiiichever “ancestral radicle” have
the same type of cardinal process. And, significant in this issue, the only
members of Leptalosia aaIioso cardinal i>rocesses are mentioned in the descrip-
tion of the species are L. truncata and L. radicans, Devonian forms. It is
just possible then that Dunbar and Condra have generalised about the
Carboniferous forms from Avork on Devonian species. We do not knoAV
AA’hether the cardinal jArocesses of the American Carboniferous Leptalosias
are of the same primith^e pattern and cannot, therefore, yet separate them
from the Scottish Carboniferous sub-genus Etlieridgina.

One other ]Aoint AAliich might be produced in support of the separate
existence of Leptalosia is the ])resence of teeth in the species of this genus,
Oehlert having defined Etlieridgina as “Avithout cardinal teeth.” The mistake
in this case is Oehlert^s, one of the co-types of E. complectens having at least
the rudiments (or A’estiges) of teeth, a condition also found among the
Carboniferous forms' of Lejjtalosia. In the face of this evidence I cannot
regard Leptalosia as other than a synonym of Etlieridgina,

Etheridgina muirwoodae n. sp.

PI. vi, figs. 14, 1 5.

1914. — Si1ro])}ialosia ,«p. Dth. fib. p. 34, pi. v, figs. 16-18.

1918 . — ‘ ^ Strophalos'ia complecfc .s” ’ Eth. fib, p. 253, pb xl, figs. 11, 12.

Syntypes. — Aus. Miis. F 16699, three specimens; Iaa'O are pedicle valves^
one sliOAA's two conjoined A'ah'es, figured by Etheridge as Strophalosia coynplect€7is
(pb 40, fig. 12). Balmaningarra, Mt. iMarmion, Kimberley Dufision. Horizon not
knoAvn.

Topotype. — Aus. Mus. F 16812, one pedicle valve, figured by Etheridge as
Strop]iaJos:ia complrcl ens (pb 40, fig. 11). Balmaningarra, Mt. Marmion. Kim-
lAorley Division. Horizon unknoAvn.

Other Material. — G.S.W.A. 10930, Mt. Marmion, Kimberley DiAusion. Hori-
zon not knoAvn. (bS.W.A. 10929, north of Barrabiddy, Xortli-West Division. Hori-
zon not knoAvn. Aus. Mus. F 3'S49S, 38499, Wandagee Station, Minilya KiA’er,
Xorth-Wc.st Division. Wandagee stage.

Diagnosis. — Shell small, transA'ersely oval to subquadrangular in outline,,
adherent by body of pedicle valve and radiating marginal sjAines. Hinge-
line less than greatest Avidth of shell, Avirh rounded alar angles.

Pedicle A’alve Avith upturned lateral and anterior margins, visceral region
adherent, flat. Hinge area Avide and fiat, psendo-deltidium large and tri-
angular, teeth strong, divergent. Strong, long, adherent spines arising from
caiMinal margin. Muscle impressions raised.

Permian Productinae and Strophalosiinae of W.A.

oo

Brachial valve concave, lamellar tending* to nodular structure. Area
and pseudo-chilidium relatively larg-e. Cardinal process inclined almost to
horizontal; muscle impressions small at base of cardinal process.

Description . — The pedicle valve is firmly affixed to the underlying shell
in most cases and hence the ornamentation is not known. In one specimen
(Aus. Mus. P 38498) the shell is geniculate and the trail is ornamented by
hne concentric lamellae and stout oblique spines. The area is wide and flat,
in length half the width of the valve and with a width one-flfth of the length.
The area is marked by longitudinal striations, interrupted medially by a
triangular deitidium.

The cardinal margin bears very long, slender spines which are cemented
to the host. These have a length much greater than the length of the shell,
one spine extending 1.3 cm. from the margin. The spines occur in a matted
group on the ears becoming more sparse towards the umbo ; they arise
perpendicular to the cardinal margin.

The teeth are strong, hollowed posteriorly to give a semi-ovoid cavity,
and diverging. The muscle impressions take a tetrahedral form, one apex
of the tetrahedron towards the umbo and beneath the area. The muscular
apophyses are slightly concave, they appear smooth and divided into a
greater and a lesser part by a narrow ridge parallel to and near their
anterior margins. The edge separating them is a thickened ridge simulating
a median septum. The anterior face of the tetrahedron is narrow, and has
■curved side edges, due to the concavity of the apophyses. Any other mark-
ings on the inside of the shell have been obliterated.

The brachial valve has the shiny lamellar structure characteristic of so
many Stroplialosias. Superimposed on these lamellae are concentric ■wrinkles
extending over the entire surface of the valve. The wrinkles are irregular
in intensity giving a general impression of a nobbly rather than a lamellai
structure. It is possible, although no evidence of their presence can be seen
on this specimen, that spines were developed over part of the visceral disc.
Marginal spines were present as shown by the numerous pustules. The
brachial valve is swollen below the umbo into a strongly convex semi-
spherical region which, undergoing rapid change of curvature, becomes flat
and so continuing across the visceral disc upturns marginally. The area of
the valve has a width about half that of the pedicle area : the chilidium is
triangular. If the valves are held in such a position that their height is
\ertical, the areas of both valves are in one horizontal plane.

Figure 9.

Cardinal Process of Etheridgina muirwoodae n. sp. (Xfi).

The cardinal process is relatively large and is inclined almost at right
angles to the area of the valves. It shows a characteristic trifld structure when
viewed ijosteriorly, although the lateral arms adhere closely to the central

K. L. Prendergast.

pillar. On each side of it are deep sockets each of which is bounded by a
curved plate arising from the side of the cardinal process and making an
angle with it of about 45°. A very short median septum is present, separat-
jiig the muscle scars which lie close to the base of the cardinal process. These
muscle impressions are not dendritic, nor do they show any division into
anterior and posterior adductors. The brachial impressions are not pre-
served. Pitting is seen over some parts of the internal surface.

The dimensions of the shells are shown by the following table: —

Co-types.

Conjoined

valves.

Pedicle valves.

Height

0*75

1-03 +

MO

0-58-f

Maximum width

0-87

1-28 +

1-41

0-92

Length of hinge-line

0-48

0-95

0-95

0-76

Variation within the species . — As may be expected in a closely adherent
species the members of this species vary widely according to the nature
of the host. The jiedicle valve may be fiat to highly convex (e.g. 10930) ; its
area, though usually horizontal, may be highly inclined. It seems, too, that
when the pedicle valve is closely adherent, it tends to assume the ornament-
ation of its host; thus, in O.S.W.A. 10930 B, where the host is a Orachial
valve of Spirifer marcoui the pedicle valves exposed are corrugated to fit in
between the striae. This feature is accentuated by tveathering.

Comparison u'ith other species . — It is with some trepidation that I have
distinguish(‘d these specimens from other species of the genus, ])ai-ticularly as
they had ah’eady been referred to E. complectens (Eth. fil.) by the author
of that species. Their characters, however, are so distinct, that I have been
forced to separate them. The specimens are larger and thicker-shelled forms-
than E. complectens \ they possess relatively large and well-developed car-
dinal areas, strong teeth and characteristic muscular iinjiression, in all these
characters being distinct from E. complectens. The mode of attachment can-
not be regarded as an important difference since, had the shells chosen a
crinoid as host, it seems probable that the spines would encircle 1he stem,
and we have as yet an insufficient number of specimens to say that they
cannot or do not choose cilnoid hosts.

The species heading of these specimens in Etheridge’s description (1918,
p. 253) is Strophalosia complectens. but elsewhere in the text and in tlf
explanation of figures he gives S. complectens. I have assumed therefore
that the title name was due to a typographical error.

It is possible that this species is really’ the adherent form of Strophalosia
etheridgei mihi. The structure of the brachial valves of the two species are
remarkably similar, the variations seen, such as the higher position of the-
muscle impressions and the more inclined cardinal proces-i in E. mii>rwooda ■
being probably modilications due to the assumption of a fixed habit. The
external a})pearances of the two species differ only in the presence of spines-
in E. muirwoodae and this too may be due to the reason noted abo-\'c.

Permian Productinae and Strophalosiinae of W.A.

0 /

VII. BIBLIOGRAPHY.

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Schmidt, H., 1929: Tierische Leitfossilien des Karbon. 107 pji., 23 pis., 20
text-figs. Lief. 6 of Gurich., G., Leitfossilien. Berlin.

Permian Productinae and Strophalosiinae of W.A,

61

Schuchert, C., 1913: Braehiopoda, |-)p. 355-420, text-figs. 526-(536, in Zittel, K.,
Textbook of Palaeontology, 2nd Edition.

8cliiicliert, C., and Cooper, G. A., 1932: Brachiopod Genera of the suborders
Ortlioidea and Pcntameroidea. Mem. Peabody Mus. Harvard, No. IV, xii ~1-
270 pp., 29 pis.

Schubert, C., and LeVene, C., 1929: Bracliiopoda, 140 pp., Pars. 42 of Pom-
peckj, J. P., Fossilium Catalogus, Berlin.

Sowerby, James, 1812: The Mineral Conchology of Great Britain. I, pp. 234 +
10, pis. 1-cii. London.

Stefanov, D. L., 1934: The Brachiopoda of the Bryozoan Limestones of the
Kolwa River Region (Northern Urals), pt. 1, Strophamenacea. Trans. Oil-
Geol. Inst., Ser. A, fase. 3.

Sutton, A. H., 1938: Taxonomy of Mississip^Dian Productidae. J. Palaeont.,-
XTI, pp. 537-569, pis. LXII-LXVI.

Thomas, I., 1914: The British Carboniferous Producti, I. Genera Pustula and
Overtonia. Mem. Geo. Sttrv. V.K., Pal. I, pp. 197-366, pis. xvii-xx, 13 text-
figs.

Tschernyschew, T. H.:

1889: Beschreibung des Central-Urals and des West-Abhanges. Allgemeino
geologische Karte von Russland, Blatt 139. Mem. Com. Geol. St.-
Petersb., Ill, No. 4, pp. viii + 393, 7 pis. (German Summary.)
1902: Die Oberearbonischeu Brachiopoden des Urals und des Timan. Lief. 1.

Text, viii + 749 pp., 85 text-figs. Russian and German. Lief 2.
Atlas, 63 pis.

Waagen, W., 1884: Salt Range Fossils, Vol. 1, Productus Limestone Fossils,
pt. 4, fase 3, Brachiopoda. Palaeont, indica, Ser XIII, pp. 611-728, pis
Iviii-lxxxi, 8 text-figs.

Wade, A., 1937 : The Geological Succession in the West Kimberley District of
Western Australia. Pep. Ansi. Ass. Adv. Sei., XXIII, pp. 93-96.

Weller, S., 1914: Mississippian Brachiopoda. Monogr. Geol. Surv. 111., 1, 508
pp., 83 pis. text-figs.

Whiteliouse, F. W., 1926: Notes on Upper Palaeozoic Marine Horizons in
Eastern and Western Australia. Pep. Aust. Ass. Adv. Sci., xviii, 1926, pp..
281-283.

132

K. L. Prexdergast.

PLATE I.

Dictyoclostus callytharrensis n, sp.

Pig. 1. Brachial valve. Syiitype. G.S.W.A. 1/4967 (b). (x §).

Fig. 2. Pedicle valve (x 1^), to show ornamentation. G.S.W.A. 1/4654.

Fig. 3, Pedicle valve. Syntype. G.S.W.A. 1/4967 (b). (x §).

Pig. 4. Lateral view of Pedicle valve. U.W\A. 12400. (x §).

Fig. 5. Brachial valve (x 11), showing hinge-line and gingliinus. G.S.W.A.
1/4654.

Fig. 6. Hinge-line (x IJ), to show ginglimus. Paratype. G.S.W.A. 1/4967
(a).

Fig.

Pedicle valve.

U.W.A. 12400. (x s).

Permian Productinae and Stropiialoshxae ok W.A.

63

PLATE I.

K. L. Fkexdkrgast.

()4

PLATK II.

Ki^'. 1. Dictyoclostus callytharrensis ii. sp. Pedicle vahe (x P';).

(i.S.W.A. 1/49(17 (a).

I'dRs. Dictycclostus callytharrensis vur. wadei. n. vai'. Ilolotvpe,

\o. 2045M. {x n).

Ilhi^e-line and iinibonal roiLiion.

3. Lateral view of liolotype.

4. I'ediele valve of lioIoty])e.

Pi^s. ~)-7, Dictyoclostus gratiosus (Waagen). Aus. AIiis. P. 37o(il

5. Pedicle valve.

(). Lateral view.

7. Brachial valve.

Pigs. S-9. Dictyoclostus spiralis (Waageii). Aus. Mas. P. 3(i->14. (x
S. Pedicle valve.

9. 1/ateral view.

Paratype

ILW.A.

(X H).

n).

Permian Peoductinae and Strophalosiinae of W.A.

G5

I’LATE II.

K. L. PRHXDEKOAST.

()()

IM.ATP HI.

Ki^'s. 1--. Linoproductus cancrinifonnis vnv. lyoni n. \ar. Ilolotvpe. Aiis.
Mils. F. 3G530. (x n).

1. Lateral view.

2. and brachial valves.

Fiy,s. 3-5. Linoproductus cora var. foordi (Mtli. fil.). O.S.W.A. 570Sb. (x i).

3. Pedicle valve.

4. Brachial valve.

5. Lateral view.

Fig. (). Waage.noconcha vagans Reed. Brachial valve. ThW.A. 20451). (x ji).

Figs. 7-1). Waagenoconcha imperfecta Ibendergast.

7. Pedicle valve. P.W.A., 20454. (x ii).

S. Brachial valve. ILW.A., 20454. (x ?i).

1). Tbnbonal region of ]te<iicle valve. P.W.A., 20457. (x H).

rLATi] III.

08

K. L. PRENDERGAvST.

PLATE IV.

Figs. ]-(>. Taeniothaerus subquadratus (Morris).

1. Braeliial vahCj exterior. XT.W.A. 12396.

2. Brachial valve, interior. IT.W.A. 21247.

3. Interior of pedicle valve. P.W.A. 4786.

4. Pedicle valve, exterior. U.W.A. 20447.

■■). Lateral view. U.W.A. 20447.

6. Hiiige-linc of crushed s]>ecinien. IhW.A. P. 21.

Figs. 7-10. Anlosteges spinosus Ilosking.

7. Pedicle valve. Ans. Alns. F. 37710.

*S. Lateral view showing recurved area. Aus. IMiis. F.

9. Brachial valve. Aus. Mu,s. F. 3S444.

10. Ih'dicle valve. Aus. Mus. F. 3S444.

Figs. 11-13. Krotovia spinulosa (-1. Howerby). Aus. Mus. 38446.

11. Lateral view.

12. Jhnlicle valve.

13. Brachial valve.

All tigures reduced to H

nat. size.

Permian Productinae and Stropiialosiinae of W.A.

G9

PLATE IV.

70

K. L. Prexdergast.

Pl.ATK \'.

Pif^s. 1:J. Stroplialosia sp. cf. Strophalosia 'beeclieri (liowU-v). I'.W.A.,
1^04-Ii). (x n).

1. Hvaeliial valve.

2. Pedicle valve.

3. Lateral view.

4. Strophalosia clarkei ( Pthei idge). Interior of hracliial valve. Aiis.
Mils. F. 3(i23o. (x

Figs. 0-12. Strophalosia etheridgei n. sp.

5 . Brachial valve in limestone. X^.W.A., 20247(a). (x H).

6. Pedicle valve (x ]}■). Syntyjie- G.SAV.A. 1/5242 (a).

7. Interior of ])edicle valve (x 1’,). Svntvpe. G.S.W.A.

1/5242 (a).

S. Lateral view (x 1,'). Syiityjie. G.S.W.A. 1/5242 (a).
0. Brachial valve (x I/.)- Hyntype. O.SAV.A. 1/5242 (a).

10. Interior of brachial vah'e (x 1\). Rvntvpe. G.SAV.A.

1/5242 (a).

11. Hiacliial vahe in limestone (x H). IWV.A., 20247 (c).

12. Pedicle valve (x ii). F.W.A., 20257.

Fig. i;}. Strophalosia gerardi King. llolotype. Fniv. (V»ll. (Jalway, I.K.S.
A^o, F.r.D. 207. Brachial valve and hinge-line, (x rs)-

Figs. 14-15. Strophalosia sp. cf. Strophalosia gerardi King, (x q)* F.W.A.
12399.

14. Pedicle valve.

15. Hinge-line of pedicle valve.

Figs. 10-17. Strophalosia gerardi King. Holotyjie. (x ii).

10. Lateral view.

17. Pedicle valve.

Figs. lS-20.

Strophalosia Jukesi Fth. fil.

LS.

Fxterual cast

of brachial valve. {:>

^ a

19.

Internal cast

of brachial valve, (x

a.

20.

Internal cast

of pedicle valve, (x

a.

U.W.A., 20200.
F.'W.A. 20204.
F.W.A. 20204.

Permian I^roduotinae and Rtrophalosttnae of W.A.

71

PLATE V.

K. L. Prkxdergast.

Figs. 1

Figs. 0

r’ig. !)
Figs, i

Fig. 1
Figs. 1

PLATE VI.

-■). Stroplialosia kimtierleyensis n. sp.

1. Brachial valve. Holotypc. P.W.A.^ 20452.

2. Pedicle valve. Ilolotype.

0. Lateral view. Ilolotype.

4. Interior of ))racliial valve. Paratypc. U.W.A., 20455.

5. Interior of pedicle valve. Para type. U.W.A., 20400.

-S. Stroplialosia multispiiiifera n. si>. nolotyi).e. P.W.A., 20458.

0. Lateral view.

7. lliiige-line.

8. I^edicle valve.

. Stroplialosia tenuispina Waagen. Pedicle val\'e. TJ.W.A., 20451.

0-12. Stroplialosia s]). A. U.W.A., 20448.

10. Interior of pedicle valve,
n. T^edicle valve.

12. Lateral view.

;i. Etheridgina complectens (Eth. fil.). Syntype. Poyal Scottish
Museiiin No. 5080.

4-15. Etheridgina muirwoodae n. sp. Syntypes. Aiis. AIus. F. 10690.

14. interior of pedicle valve and exterior of brachial valve.

15. Interior of pedicle valve.

All figures reduced to n nat. size.

Pp:Ri[IAN PRODUCTIXAE AXD StROPHALOSIIXAE OF W.A.

By Autlioi-ity; ROHicirr U. MiLr.KU, Government Printer, Perth.

The Occtjbkence or the Genus Conoclypus in the North-West 75

Division, W.A.

2.— THE OCCURRENCE OF THE GENUS
CONOCLYPUS IN THE NORTH-WEST DIVISION,
WESTERN AUSTRALIA.

By Irene Crespin, B.A., Commonwealth Palaeontologist..

Read 9th September, 1941 ; Published 21st May, 1943^

Communicated by Professor E, de C. Clarke.

In 1939, Mr. E. A. Rndd, M.Sc., when associated with Oil Search, Ltd.,
of Sydney, collected three specimens of a large echinoid from the east flank
Cape Range, Exmoiith Gulf Station, North-west Australia, whilst further
examples were recently discovered in the same region by geologists attached
to Caltex (Australia) Oil Development Pty., Ltd. All specimens have kindly
been made available for examination. The tests are embedded in a pink to
ochreous limestone containing species of Lepidocyclinae characteristic of the
Middle Miocene in the Indo-Pacific. The form has been determined as Cono-
clypus, a rare genus only previously recorded from the Eocene in areas outside
Australia.

The occurrence of Conoclypus, according to Morley Davies (1935) is
“ widespread in the Cretaceous, but unknown in Cainozoic of America or
Australia.” It is recorded from the Eocene of India, Egypt, Madagascar,
Mozambique, and Persia, as well as from countries in southern Europe. “ Its
geological range thus follows the large Nummulites and like them, it never
reached America and is apparently extinct after Middle Eocene.”

All specimens are in the Commonwealth ‘Palaeontological Collection ^
Canberra, Australia. The photographs of the types were taken by Dr. F. W.
Clements, Australian Institute of Anatomy, Canberra.

DESCRIPTION OF SPECIES.

Plate I., figs 1—3.

Class ECHINOIDEA Brown.

Order HOLECTYPOIDA.

Family ECHINONEIDAE.

Genus CONOCLYPUS Agassiz.

Conoclypus westraliensis sp. nov.

Holotype . — Although a specimen has been selected as a holotype, the
species cannot be fully described without referring to characters which are
better illustrated in further specimens.

Test moderately large, almost circular in outline, broadest beliind apical
system above periproct ; moderately convex to subconical on aboral surface,,
with a flat adoral face. Entire test covered with small tubercules sunken
in rounded scrobicules. Apical system slightly praecentral or forward.
Ambulacral and interambulacral plates only visible in worn specimens.
Genital plate distinct with four genital pores. The two pore fields, which
outline the five petals, are petaloid in shape. The pore fields, wliich are com-
posed of pore pairs of the dot and dash type, in wLich one pore becomes slit-

76

Irene Crespin.

like, are wide, extending from apex to ambitus, and converging towards apex
and peristome. Peristome on adoral surface, central and pentagonal, with
five interambulacral bourrelets, representing the rounded blunt projections
from each ambulacrum adjacent to the peristome, and five ambulacral
phyllodes, representing the ambiilacral pores. The phyllodes indicate tlie
petaloid arrangement developed on the aboral surface, and are distinct on one
specimen. The five bourrelets and five phyllodes form the floscelle. The
periproct is on adoral surface at posterior end of test in the fifth interambulacral
area close to ambitus.

Dhne'nsio7is. — Height — Holotype, 49 mm. ; Paratype, 41*5 mm. ; Plesio-
types, (1) 41 mm., (2) 41*5 mm. Diameter of adoral surface from periproct
to 3rd ambulacral— Holotype, 74 mm. ; Paratype, 70*5 mm. ; Plesiotypes,
(1) 61 mm., (2) 66 mm.

Occurrence. — Holotype, Gorge east flank of Cape range, west of Exmouth
Gulf Outcamp, North-west Division, Western Australia, coll. E. A. Rudd.
(Com. Pal. Coll. No. 184). Paratype, same locality as holotype. (Com. Pal.
Coll. No. 185). Plesiotypes, Open gorge, north of Mt. King, Cape range, coll.
E. A. Rudd ; 4-7 miles from mouth of Badjirrajirra Creek, Exmouth Gulf,

coll. Caltex (Australia) Oil Development Pty., Ltd. (Com. Pal. Coll. Nos.
186, 187).

Ohservatioyis.—Morlej Davies states that “ the genus is remarkable in
having jaws of a Holectypoid, tlie Cassidulinoifl floscelle and Clypeastroid
petals.” The genus is typically represented in C. westralicnsis, the four
specimens availal:)le for examination being uniform in shape but varying in
height and diameter. Tate, in 1893, describetl from the Tertiary beds at
Table Capo, Tasmania, an ecliinoid under the name of Conoclypens rostratus.
The specimen, which is in the Tate Museum, Geology Department, University
of Adelaide, has beoai re-examined, and belongs to the genus Echino layy^jnis.
At the time wlien Tate describcnl his form, the beds at Table Cape were con-
sidered Eocene in age, but they are now referretl to the Miocene, being most
probably Middle Miocene. Tlie limestone in which C. westraliensis is found,
contains a foraminiferal assemblage typical of the Middle Miocene in the
Indo-Pacific region and includes such forms as C ycloclypcus posteidae Tan
and Lepidocycliyia (Tryhliolepidina) ynartini (Schlumberger). There is, there-
fore, little doubt that the genus ranges to a much liigher horizon in the Tertiary
in the Tndo-Pacific region than previously considered.

REFERENCES.

Agassiz, L., and Desor, E., 1847. Catalogue raisonne des families, genres, et des especes
de la classe des echinides. Ann,. Sci. Nat. (3), Zool. 8, p. 167.

Davies, Morley A., 1934-1935. Tertiary Faunas. Vol. X.. II.

Tate, R.. 1893. Unrecorded Genera of the Older Tertiary Fauna. Journ. Hoy. Soc. N.S.W
27, pp. 167-197.

Wright, T., 1857. A Monograph on the Britisli Fossil Eehinoderinata from the Oolitic
Formations. Part III. Pal. Soc. London, 9.

DESCRIPTION OF PLATE.

All figures f natural size.

Figs. 1-3. Conoclypus westraliensis sp. nov., Cape Range, Exmouth Gulf Station, North
west Division, Western Australia.

,, 1. Side view of Holotype, showing typical Conoclypus shape and characteristi

surface ornamentation.

„ 2. Adoral surface of paratype, showing peristome, periproct, 5 interambulacr^

bourrelets and 5 ambulacral phyllodes,

„ 3. Aboral surface of holotype, showing 4 genital pores and slightly praecentral

apical system.

.

a

The Occurrence of the Genus Conoclypus in the North-West 77

Division, W.A.

"v

--

mm

mm

tK'V

ih.

S^N:. .’ '

K"'-.

'Xm : •:

r\

J
. v>

• c

Fibrofebrite and Copiapite from Yetar Spring, near Chidlows. 79

3 .— fibroferrite and copiapite from yetar
SPRING, NEAR CHIDLOWS.

By Rex T. Prtder, Ph.D., B.Sc., F.G.S.

Read 14th October, 1941 : Published 21st May, 1943.

Yetar Spring (Lat. 31° oG*o^ S., Long. 11G° 21-4' E.) is situated in the
upper reaches of the Helena River some 51 miles S.S.E. from the junction
of the main hTortham and York roads {see W.A. Govt. Litho. 2 /80).

My attention was drawn to this locality by Mr. L. Burgess of Fauldings,
Ltd., Perth, who collected encrusting material from the mound built up by
this spring. This material, which Mr. Burgess gave to me for determination,
proved to consist largely of a mixture of fibroferrite, copiapite, and melanter-
ite. As this occurrence of fibroferrite was the first record of this mineral for
Western Australia the locality was visited by Professor Clarke and myself
in April of this year, and a closer examination of the occurrence was made.

THE MOUND SPRING.

The spring, situated on flat ground, several hundred yards south of the
Helena, has built up a mound 1|- chains in diameter and about 15 feet high
(fig. 1). The mound consists of a black peaty earth which is a greyish colour
when dried. This material, on ignition, yields a greyish diatomaceous earth
consisting entirely of diatom remains and delicate rods of amorphous silica.
The latter are broken and do not show any pointed ends or a central canal
and are therefore not sponge spicules — they may be silicified plant fibres.
A partial analysis of this diatomaceous earth (dried at 100°C.) gave ; —

0/

/o

Loss on Ignition” ...

21-85

f^iCb

69-71

FcoOs

3-46

Rest ...

4-98

100-00

Anal. : R. T. Prider.

According to tlie owners of this property, the mound from which the spring
issues has gradually increased in size, apparently by growth from below,
during the forty years that it has been known, so that several enclosing fences
have been destroyed. About LJ- chains wnst of the main spring there is another
small spring and the ground surface over an area of about ten scpiare yards
is slightly uplifted.

The water from the spring is rich in solids (determined at 0'3077 parts
per cent.) and qualitative tests indicate that chlorides are the dominant
constituents with minor amounts of sulphates. The metallic ions present are
mainly alkalies with lesser amounts of magnesium. No iron is present in
the water, although in the channel through which the spring water runs over
the surface of the mound there is a copious precipitate of flocculent iron oxide.

The sample collected by Mr. Burgess formed an encrustation on the peaty
diatomite in the vicinity of the channel draining the water from the top of the

80

Rex T. Prider.

mound. It consisted of a mixture of white fibroferrite (predominant), yellow
copiapite, and colourless melanterite. None of this material remained at
the time of my visit in April, 1941 (possibly having been dissolved by rain,
as these minerals are all w’atcr-soluble) and the only ferric sulphate minerals
seen w'ere from encrustations on the surface of small heaps of spoil from shallow
holes which had been dug on the surface of the mound — such material con-
sisted largely of copiapite and melanterite with no libroferrite. Several
samples were taken from the encrustations: ( 1 ) of moist yellow material,

and ( 2 ) of dry earthy yellow material.

Figure 1.

General view of the niouiul at Yetar Spring. The figure on right

gives tlie scale.

The f(‘rric sulphate minerals are : — •

A. Fibroferrite . — Present only in the sample collected by Mr. Burgess.
It was identified optically and the determination checked by qualitative
chemical tests. The properties of this mineral are : —

White in colour in the mass and soluble in cold water, the resultant
solution giving positive reactions for Fe"' and SO 3 only. Yields water
and then SO 2 in the closed tube. Under the microscope it is seen to be
in tiny colourless acicular forms with Z [| length, y = 1*600 + *002 and
oc = 1 • 553 + * 002. These refractive indices are somewhat higher than
those given by Larsen and Bennan (1934, p. 103) for fibroferrite, viz.,
y ~ 1*575 and oc 1*533.

None of the Yetar Spring fibroferrite could be obtained free from copiapite
and no quantitative analysis has been made. The small lumps of white
fibroferrite (about J inch diametei’) when broken are seen to contain a central
core of copiapite.

B. Mekmterite was identified optically from the samples collected by both
Mr. Burgess and myself. My samples consisted entirely of copiapite and
melanterite. In the damp material (sample 1) melanterite was more abundant
than in the dry sample (No. 2) which consisted predominantly of copiapite
with lesser amounts of melanterite. An estimation of ferrous iron m the
moist material after drying in a desiccator over calcium chloride gave FeO

Fibroferrite and Copiapite from Yetar Spring, near Chidlows. 81

7 • 38 per cent, (as compared with FeO 4 * 50 per cent, in the naturally dry sample
No. 2 — see Table 1 below). The melanterite occurs as colourless weakly bi-
refringent granules with refractive index 1 • 48.

C. Copiapite is the most abundant constituent of the naturally dried
material (sample 2). It is earthy yellow material, soluble in cold water, yield-
ing a brown acid solution. Under the microscope it is seen to be in micro-
scopic yellow non pleochroic plates with high birefringence and y = 1-578.

An analysis was made of the copiapite-rich sample (No. 2) and the result
is shown in Table 1.

Table 1.

Analysis of Copiapite-rir.h Material {Sample 2).

Analysis on material dried in desiccator over calcium chloride for seven days.

Insol. (1) ...

•

/o

0*88

Mol.

prop.

Melanterite.

Copiapite.

No. of Metal
Atoms on basis
of 6 S.

FeO

4-50

•063

■063

, . •

Fe/J, ...

21 -39

-134

•134

3-78

AI 0 O 3

Nil

, • •

Cab

Tr.

MgO

1*27

■032

■032

0-45

K 2 O

0-25

-002

•002

0-06

Na.^O

2-17

■035

•035

0-99

SO,

39-17

•490

■063

•427

6-00

Hob( 2 ) ...

30-37

1-687

-441

1*246

35-10

Of

Tr.

...

. • .

• • .

. . .

100-00

Anal. • R. T. Prider.

(^) Insoluble in cold water.

(^) By difference. A separate determination on another sample gave “ loss on ig-
nition ” 65*82 per cent.

On the assumption that all of the FeO is in the form of melanterite the
residue (copiapite) has the formula (Na 2 , Mg)o. 97 (OH)i. 94 .

- 00 -

16-58 H 2 O, which agrees fairly clo.scly with the formula X( 0 H) 2 .Fe"' 4 (S 04 )g.
n H 2 O suggested by Peacock (see Berry, 1939, p. 182). Following the nomen-
clature proposed by Berry (1939) this mineral would be a soda-magnesio-
copiapite.

THE ORIGIN OF THE MOUND AND THE ASSOCIATED FERRIC

SULPHATE MINERALS.

The mound surrounding the spring has apparently grown by addition of
material from below. It ajopears most probable that the ferric sulphate
minerals have been derived from a mass of pyritic material lying below the
surface. The surroimding country (although outcrops are scarce in the
imnu'diate vicinity of the spring) is made up of Pre-Cambrian granitic rocks
which would not be expected to contain any appreciable amount of pyrite —
nevertheless pyritic masses are known to occur in the Darling Range granite,
for example, at the Canning Dam quarry I have observed a well developed vein,
several feet wide, of crushed granite heavily mineralised with pyrite. Again
there may be a dolerite dyke carrying pyrites below the mound spring. Material
<lerived from such a mass of decomposing pyrite lias evidently been carried to
the surface by the spring waters and deposited in the porous diatomite of the

82

Rex T. Prideb.

mouncL The ferric sulphate minerals crystallising throughout the j^orous
diatomite would probably supply sufficient force to cause the slow uplift of
the mound that has been observed.

ACKNOWLEDGMENTS.

I wish to acknowledge my thanks to Mr. L. Burgess for drawing my
attention to this deposit. The expenses entailed by a visit to the locality
were met by a Commonwealth Research grant for work on the Pre-Cambrian
rocks of W.A. My thanks are due also to Professor E. de C. Clarke for his
assistance in connection with the visit to the locality and during the pre-
paration of this paper.

REFERENCES.

Berry. L. G., 1939. “ Composition and Optics of Copiapite ” (abstract). Amer.

Vol. 24, p. 182.

Larsen, E. S. and Berman, H., 1934. “ The Microscopic Determination of the Non-opaque

Minerals, 2nd Ed.” U.S.G.S. Bull. 848.

Geology and Petrology of Part of Toodyay District, W.A. 83

4 .— THE GEOLOGY AND PETROLOGY OF PART

OF THE TOODYAY DISTRICT,
WESTERN AUSTRALIA.*

By Rex T. Prider, Ph.D., B.Sc,, F.G.8.

(Df'pavtment of Geology, University of Western Anstraiia).

R('ad 12tb August, 1941 : Published 14th March, 1944.

(’ONTENTS.

Page

T. — Introdvcuton 84

II. — Geological Structure of the Area ... ... ... ... ... ••• 85

ITT. — Petrology ... ... ... ... ... ••• ••• .•• ••• ••• 88

A. — The Jimperding Series ... ... ... ... ••• ... 88

( 1 ) Quartzites ... ... ... ... ... ... ■ ■ ■ ■ - ■ 88

(а) Upper quartzite ... ... ... ... ... ••• 90

(б) The lower quartzites ... ... ... ... ... 91

(c) The origin of the quartzites ... ... ... ... 92

(^) The fabric of the quartzites ... ... ... ... 94

(2) Calc-silicate rocks ... ... ... ... ... .•• ... 94

(3) Mica schists ... ... ... ... ... ■■■ .■• ••• 97

(а) Sillimanite schists ... ... ... ... ... ... 98

{h) Garnet-plagioclase-biotite schist ... ... 99

{r.) Gedrite-cummingtonite-plagioclase schist ... ... ... 100

{d) Andalusite schists ... ... ... ... ... ... 101

(4) Hornblende schists (plagioclase amphibolites) ... ... ... 104

(5) The Granite Gneisses and the associated xenoliths ... ... 107

{a) Upper Granitic Gneiss ... ... ... ... ... 108

(i) Biotite-microcline granite gneiss ... ... ... 108

(ii) Microcline granite gneiss ... ... ... ... HI

(hi) Biotite-oligoclase granite gneiss ... ... ... Ill

(б) Xenoliths in the Upper Granitic Gneiss Ill

(i) {Schistose plagioclase amphibolites ... ... ... Ill

(ii) HornVdendic schlieren... ... ... ... ... 11.2

(hi) Biotite granulite xenoliths ... ... ... ... 112

(iv) Eulysitic rocks ... ... ... ... ... 113

(c) Conclusions regarding the origin of the Upper Gneiss ... 114

(^^) Lower Granitic Gneisses ... ... ... ... ... 114

(c) Xenoliths in the Lower Granitic Gneiss ... ... ... 115

(i) Eulysitic rocks ... ... ... ... ... 115

(ii) Calc-silicate rocks ... ... ... ... ... 118

(iii) Cordiei’ite-anthophyllite rocks and related types ... 118

(iv) Amphibolite xenoliths ... ... ... ... 119

(f) Hybridisation of the Lower Granitic Gneiss ... ... 122

Ig) Conclusions regarding the origin of the Lower Gneiss and

the associated xenoliths ... ... ... ... ... 123

B. — The Younger Igneous Inirusives ... ... ... ... ... ... 124

(1) The granites 124

(а) The normal granites ... ... ... ... ... ... 124

(б) Garnet-muscovite granites ... ... ... ... ... 124

(c) Pegmatites, garnet aplites, and later quartz veins ... 124

(2) The greenstones ... ... ... ... ... ... ... 125

(а) Quartz dolerites ... ... ... ... ... ... 125

(б) Ultrabasic intrusives ... ... ... ... ... ... 128

(i) Metamorphosed ultrabasic sills ... ... ... 128

(ii) Serpentines ... ... ... ... ... ... 128

IV'. — Economic Geology 129

(1) Gold 129

(2) Refractories ... ... ... ... ... ... ... ... 129

(a) Sillimanite ... ... ... ... ... ... ... 129

{b) Andalusite ... ... ... ... ... ... ... 129

V. — Summary ano Conclusions ... ... ... ... ... ... ... 129

VI, — Acknowledgments ... ... ... ... ... ... ... ... 131

Vll. — L lst of References 132

Western Australia.

k 30/48.

84

Rex T. Prider.

I. INTRODUCTION.

The area which measures about 12 miles in an East-West direction, by
three to four miles in a North-South direction, lies several miles to the south
of the town of Toodyay, which is situated about 50 miles north-east from
Perth, It is occupied almost entirely by a series of crystalline schists to which
the name “ Jimperding Series” has been given (Clarke, 1930, p. 167) and
which is believed to be of an early Pre-Cambrian age. This series comj^rises
politic and psammitic metasediments with intercalated basic and acid igneous
bands. A study of the politic members shows that over the wliole area mapped
the rocks lie within the sillimanite zone.

This series has been intruded by granite (and its associated aplitic and jjeg-
matitic dykes) and afterwards by a series of quartz dolerite and rare ultrabasic
dykes — none of these igneous intrusions has metamorphosed tlie older rocks.

The only rocks of later age than the quartz dolerite series are of supei--
ficial character — laterites (duricrust, Woolnough, 1930) and recent alluvial
deposits. The duricrust occurs at an elevation of about 900 feet above sea-
level, and has been developed over all the older rocks, even the extremely sili-
ceous quartzites. Neither it nor the alluvial dejDosits will be considered in this
paper, which deals with the Pre-Cambrian rocks.

A brief accoimt of the rocks from the western part of this area has betai
published (Prider, 1934, pp. 1-16) but, ’other than this, there is no account of
the petrology of any of these Pre-Cambrian rocks. Simpson has described a
number of the minerals occurring in the metamorphic rocks of the Darling
Range, several from the area at present under consideration, viz., andalusite
from Jimperding (Simpson, 1928, p. 50), sillimanite, also from Jimperdinir
(Simpson, 1936, p. 10), and grossularite from Key Farm (Simpson, 1937, p. 32 j.

Clarke (1930, p. 167) shows that the Jimperding Series forms the greater
part of the northerly extension of the Darling Range. From the vicinity of
Toodyay, the rocks extend in a belt which has been traced as far north as the
Irwin River District (about 200 miles north from Toodyay) and Ninghanboun
Hills (Simpson, 1931, p. 138). I have examined the rocks in the Irwin River
District and they appear to be essentially the same as those developed at
Toodyay.

To the south of Toodyay the Jimperding Series has been traced through
Clackline (the series exposed here is the upper portion of the Jimperding
Series) to York, a distance of 30 miles.

Previous to the publication of a paper by Forman (1937, pp. xvii.-xxvii.),
the Jimperding Series had been regarded as the oldest formation in the Western
Australian Pre-Cambrian shield, Forman (1937, p. xxv.) regards the green-
stones ” of the Kalgoorlie Series as being older than the Jimperding (= Yil-
garn) Series. Feldtmann (1919) describes a greenstone series at Bolgart (situ-
ated approximately 20 miles north from Toodyay) and correlates it with the
Kalgoorlie Greenstone Series and Forman considers that these rocks probably
underlie the Jimperding Series (1937, p. xxvi.). I have examined slices of some
of the Bolgart rocks collected by Mr. R. W. Fletcher and find, in them, the
counterpart of rocks occurring as xenoliths in the granitic gneisses in the
Toodyay District — this would be confirmatory evidence that the Jimperding
Series is younger than the Bolgart “ greenstones,” were it not that similar
rocks are interbedded with the metasediments of the Jimperding Series.

Geology and Petrology of Part of Toodyay District, W.A. 85

This paper has been subdivided into two main sections, thus
1. The Geological Structure.

Tl. Petrology —

(A) The Jimperding Series.

(B) The later igneous intrusions.

In view of the variety of rock types developed in the area, theories of
origin of each main group are discussed immediately after the description
of typical members of that group and these theories are briefly recapitulated
at tlie end of the paper.

II. THE GEOLOGICAL STRUCTURE OF THE AREA.

The regional strike throughout the Pre-Cambrian rocks in South-Western
Australia is N.W.-S.E. In the area under consideration, there is a distinct
departure from this N.W. regional strike, for, over a great part, the strike is
more nearly E.-W.

A Pre-Cambrian metasedimentary series (the Chittering Series) has been
described by Miles (1938) from the Chittering Valley, some 15 miles west from
the Toodyay area. The dominant strike of these rocks is N. to N.W. These
rocks are similar in character to the Toodyay rocks, but no information regard-
ing the relationship of the Chittering and Jimperding Series is available, as
the country between Toodyay and tlie Chittering Valley has never been
geologically examined.

Soutli of tlie Toodyay area, the Jimperding Series has been noted at
Clackline and York and in both places the dominant strike is in a northerly direc-
tion.

It will be seen, therefore, that the Toodyay area is one of abnormal strike
- -unfortunately, field work has not covered a sufficient area to warrant any
dthnite conclusions being drawn as to the true character of the structure,
l)iit in tlie following pages the structure is described so far as it is known.

From tlie geological map of the area (Plate I. ), it will be seen that the whole
are'a, except tlie S.W. corner, is occupied by a confoianable series (note, how-
(‘^■er, that at a position about 300 chains W. and 80 chains N. of datum, the
out(.‘r band of granitic gneiss appears to transgress the quartzite-hornblende
s('hist-mica schist bands). The main structural features can be seen best in
the accompanying sketch map (figure 1), which has been simplified from
th<‘ more detailed, larger-scale, maj).

Two wide bands of granite gneiss are interbedded v'ith the metasediments
-the banding everywhere conforming to that of the metasediments (except
in the one place noted above). In the south-west part of the area, the bound-
ai’y between granite gneiss and metasediment on the western side strikes nearly
E.-W., but farther east the strike swings round to a south-easterly direction.
The northern Iioundary of this upper gneiss band does not run parallel to the
southern boundary, except in the westernmost part of the area. Towards
the centre of the area it swings to the N.E. for a distance of approximately
thre(^ miles and then turns sharply to a S.S.E. trend. The outer granitic
gneiss (termed “ Lower Gneiss ” in figure 1) runs more or less parallel to the

86

Rex T. Prider.

1

i-

Fig. 1.

Structural Sketch Map of the Toodyay Area.

Geology and Petrology of Part of Toodyay District, W.A. 87

northern boundary of the upper granite gneiss and is bounded on both sides
by metasediments, the dip and strike of which conform to that of tlie gneiss.

Regarding the dip of these rocks — in the western half of the area the dip
is constantly south at amounts ranging from 15° to 40° ; in the centre of the
area the general dip is at fairly flat angles to the south-east but there is a con-
siderable amount of minor folding in the metasediments in this part ; at the
place where the strike suddenly changes to the S.S.E. in the N.E. part of the
area the dips are very steep to the south, and thereafter fairly steep to the
east.

From the fleld mapping the. structure of this area is interpreted as : —

A major anticlinal structure (sec flgure 1), the axis of which traverses
the centre of the area in a direction striking approximately N.N.W. and
pitching to the S.S.E., which has, on its eastern limb, a recumbent syncline
with its axial plane striking N.N.W. and dipping towards the east at approxi-
mately 60°. The axis of this fold also pitches to the south. The sequence as
it appears in the eastern part of the area is therefore inverted. These struc-
tures are illustrated on the cross sections appearing with the geological map
in Plato 1.

This idea of the structure of tlie area Y^ould receive considerable support
if it could be proved that the beds of the north and north-east parts of the area
were older than those of the south-western parts. Up to the present, however,
there is very little evidence othei' tlian that afforded by occasional drag folds,
for determining the age relations of these rocks. Complete recrystallisation
has almost obliterated graded bedding and similar structures which might
have afforded criteria for determining the stratigraphical succession. (Some
obscure current bedding structures have been noted but no certain interpreta-
tion of these was possible). All the rocks are in the sillimanite zone, and
consequently grade of metamorphism affords no information as to the succes-
sion. Read (1936, p. 473) has pointed out that the abundance of andalusite
in narrow politic bands may be due to graded bedding — -the lower gritty pos-
tions of the band being almost free from andalusite, which becomes more
abundant in the upper more politic part of the band, and he has applied this
method to determining the stratigraphical succession in the Dalradian rocks
at Banff. In the area at present under discussion this variation in size and
abundance of andalusite in the upper politic rocks of the series is not present.
As far as can be seen the rocks throughout this band are fairly uniform in
composition. There is, as will be shown later, a constant variation in the
development of the andalusite in this band, but this extends over a consider-
able wddth and is not due to any variation in the argillaceous components of
the original sediment, but rather to a variation in the temperature conditions
to which the rocks have been subjected.

Drag folded structures in the Cjuartzites at a position 140 chains east
and 225 chains south of datum indicate that the easterly dipping metasediments
at that locality are on the western limb of an anticline overturned to the west
and that the se([uence in this vicinity in inverted.

If, as suggested by the drag folds at 140 E., 225 S., the rocks of the north-
ern and eastern part of the area are older than those to the south-west (if they
are not, the whole succession over the greater part of the area surveyed must be

88

Kex T. Prider.

inverted), then the order of succession, with approximate thickness of the

individual beds

is as follows : —

Upper ....

Andalusite-muscovite schist ....

> 250'

Quartzite

Upper Granite gneiss (with rare basic igneous

and

500'

meta-sedimentary xenoliths)

1900'

Quartzite

110'

Hornblende schist

40'

Quartzite

570'

Sillimanite schist

100'

Quartzite

375'

Lowei’ Granite gneiss (characterised by the j)resence

of basic igneous and sedimentary xenoliths)

5,400'

Quartzite

050'

Hornblende schist

35'

Lower ....

Quartzite

Unknown

These rocks form the Jimperding series and are intruded in succession
by the following -

1. Coarse non-foliated microcline granite.

2. Pegmatites, garnet aplites, and garnet muscovite granites.

3. Quartz dolerites.

4. Serpentinised ultrabasic sills and dykes.

The age relation of the quartz dolerite series and ultrabasics is not at
present known.

Large scale faulting in the metamorphic rocks has been noted in several
places, the most notable being in the part of the area situated about one and
a-half miles east from Nardi Trig Station where a faulted block of the meta-
morphics is found. This block has suffered a displacement of approximately
one and a-half miles along the fault plane (see E.~W. section through datum
on Plate I.).

The lithology and structure of the area are reflected in the topography.
The metasedimentary areas are all rough country, whereas the granite gneiss
and granite areas are comparatively smooth. The main stream, the Avon
River, as will be seen from the geological map (Plate T.) follows the structure
very closely, and its main tributary (the Jimperding Brook) behaves in a
similai’ fashion in the upper part of the series.

The distribution of the later quartz dolerite dykes is interesting in con-
nection with the structure of the area. A glance at the geological map shows
that they trend generally in a N.N.W.-S.S.E. direction. There are some
exceptions to this, but the greater number of dykes have this trend. They
have apparently come up along lines of weakness approximately parallel to
the tectonic ‘‘ strike " (or tectonic axis).

III. PETROLOGY.

A . — The Jimperdinq Series.

Tlie metasedimentary members of this series, \dz., the (juartzites and
mica schists will be described first and the igneous members (plagioclase
amphibolites and granite gneisses) later.

(1) The Quartzit-s.

These are mostly coarse grained alnmst pure ([uartz rocks occurring in
well defined bands. In the field they have a well bedded, flaggy appearance
and are much iointed.

Geology and Petrology of Part of Toodyay District, W.A. 89

On all the bedding surfaces a well marked lineation (figure 2), due to
elongation of mica flakes, is visible. This lineation is often accompanied by
a corrugation of the surfaces of the Cjuartzite flags. Unfortunately the sig-
nificance of this lineation as pointed out by Phillips (1937, p. 591) was not
appreciated at the time when the original field survey (made j^rior to 1936)
was being carried out, but certain significant localities within the area have
since been re-examined and the direction and pitch of these lineations noted.
The light that these observations have thrown on the geological structure will
be given in a later section.

Pig. 2.

Quartzite outcrop in Gorge Creek, near Key Parni, Toodyay, show-
ing well marked h-lineations. The clinometer indicates the strike
and rocks dip towards the observer in the direction of the arrow.
The hammer handle lies in the direction of the b-lineations which
pitch to the left-hand side of the outcrop. The well-marked joint-
ing of the quartzites is a noticeable feature. (Eeproduced from
^‘Junior Geology by E. de C. Clarke and L. P. Hanrahan, by kind
permission of "the University of Western Australia Textbooks
Board.)

The structure in all tlie specimens examined microscopically, is similar,
viz., coarsely granoblastic consisting almost entirely of irregular interlocking
grains of cpiartz. All signs of clastic structure have been completely obliterated
by recrystallisation. In all the rocks examined there is slight undulatory ex-
tinction, but no other signs of crushing in the cpiartz. A pale greenish chrome
muscovite is the most common constituent other than rpiartz, and is present
in all the specimens examined. Occasionally narrow seams up to 2 inches
thick of this mica are interbedded with the quartzites. Felspar (mainly
microcline with a little oligoclase) is a common constituent, but rarely exceeds
5 per cent of the rock. Minor minerals present in varying amount, are mag-
netite, apatite, rounded pink zircons, biotite, and rutile. These heavy ”
minerals are occasionally concentrated into bands (figure 4B).

In the earlier examination of these rocks from the Jimperding Area, it
was thought possible to distinguish the various quartzite bands by means of
the mmor constituents (Prider 1933, p. 7). Extension of the survey and an
examination of a greater number of the lower quartzites has indicated that.

90

Rex T. Pkider.

while possible to distinguish between the upj)ermost horizon and the bands
in the lower part of the series, these low er members could not l^e distinguished
from one another.

In this paper, therefore, the cpiartzites will be considered in two main
groups —

(u) The u]jper (piartzit(' (Xo, 5 fpiartzite of the earlie?' paper).

(6) The lower quartzite horizons.

(u) The Upper Quartzite is characttalsed by the mode of occurrence of
ehrome muscovite and almost complete absen(*e of felspar. These rocks are
all even, coarse-grained types, consisting almost wholly of (juartz, in which
the average grain size is of the order of 3 mm. diameter, although much
coarser vaiieties are often encountered. The remarkable evemaess and coarse-
ness of grain indicate the high degree of metamorphism to which these rocks
liave been subjected (Harker, 1932, ]>. 97).

A. Micaceous quartzite. Section ])erpendiculnr to bedding.
Shows the coai’Se granoblastic structure and parallel oriented
)‘ods of chrome-inuscovitc.

B. Micaceous quartzite. Wection ])aranel to bedding showing
tendency of idioblastic mica to be elongated in a common direc-
tion ■ direction of lineation seen in hand specimen.)

Under the microscope' the irregular interlocking cjuartz grains frecpiently
exhibit a marked elongation in sections cut normal to the bedding. Undulose
extinction often accompanied by incipient cracking is a common feature.
Minute dusty inclusions, many of w’hich are gas-liquid inclusions, are alw'ays
present and appear to be arranged in lines normal to the direction of elongation
of the grains (figure 3A), These appear to be directions of tension joints
(“ ac joints”) as described by Fairbairn (1937, p. 89), although this cannot
be verified until fabric analyses of these rocks are made.

The presence of abundant chrome muscovite inclusions in the quartz
grains is the most remarkable feature of the upper quartzites. These are
all arranged in parallel orientation as .shown in figures 3 (A) and (B). In

Geology and Petrology of Part of Toodyay District, W.A. 01

sections normal to the bedding the mica appears in innumerable minute paralh'l
rods. Sections parallel to the bedding (figure 3B) show tliat the minute mica
flakes are all euhedral and the marked lineation noticed in thi' hand specimi'n
becomes evident under the microscope in the elongation of tlu^se flakes in
a common direction.

The mica, in hand specimen, is a pale bluish-grein colour, but under
the microscope it appears colourless, except in thicker sections, when it has
a bright light greenish colour with pleochroism -

X pale bluish ; Y Z pale yellow-green ; absorption X ^ Y Z.

^ Dti06 ; ( -)2V 31°~33° ; Dispersion distinct r >

A determination of the Cr .>03 content of mica isolated from a narrow
band (2 inches wide) of chrome mica in ojk' of the Lower Quartzites gave —

0-22 p(T cent.

This is considerably lower than the CT. 2 O 3 of tlu‘ normal clirome muscov'ites
cjuoted by Do('lt('r (1917, p. 42S). Of the six analys(‘s (|uot(‘d, the lowest,
^ is 0-S7 p(T cent, and it goes as high as 3*95 per cent in the original

fuchsite from Zillerthal. Hutton (1940, p. 330H) has di'scribi'd a chromic
musco\'it(' (H)iitaining 0-27 ])('r c('nt Gr .203 and has not(‘d liow a v(‘ry

small chromic contcait is sufli('i(‘nt to )>ro(luce a bright gr(H 3 i colour in a thick
flake of the miiK'ral.

Partridge (1937, p. 457) has recently describt'd a similar mica from Maslii-
shimala, Transvaal. It (‘arries 0-S5 p(‘r cint C'ro 03 and Partridge considc'rs
that chrome muscoviti' is a bi'tter name for this mim'ral than fuchsite as it
was })r(“viously termed. In its o])tics : Pleochroism X blue, yellowish

p-eim, Z bright gret'ii (bluish), )3 = 1-590, (-) 2V 35° : it agrei's ciosi'ly
with the mica in th(' Toodyay (juartzites. The (-hrome muscovite, ai^cording
to Partridges occurs (juite commonly in the <{uartzit(>s of tin* Swaziland Systean
in South Africa and it is tlu're'fore similar in occurrence' to the' e-hrome'-muscovite
in the Toenlyay e|iiartzite*.s.

The' source of the chromium, whie-li is so wiel(*ly disse'ininate'el througii
these ejuartzite^s, is unknown -no chrome-be'aring detritals ha\e' be'cn nott'el
in those rocks anel ne) ultrabasic re^cks (e)the'r than se'\ e‘ral small beielies youngea*
than the (juartzite's) are known in the area. Partrielge' (1937, ]). 459) has
noteel the' [ire'sence of traces of chromium in minerals e-onnecte'el with, and
ceintaineel in, the granites e>f North-Last Trans\'aal, anel alse> that the oce-ur-
reaiea's of chrome-musceivite are' conne'e'te'el with the' granite inti’usions anel are>
founel einly in close proximity tei the' gi-anite'- -he cone-luele's, the'refore, that
the e‘hrome-musce)vite is gene'tie*ally relateel to the granite'.

An intere'sting assoe*iation of teiurmaline' with the e-hronu' muscovite was
note'el in the' ejuartzite\s in the Gorge at Kew Farm -hea-o a fe*w platy crystals
e)f tourmaline u]i to 2 cm. long, coateel with the' gre'e'nish mica, we're founel
eai the be'deling surface's of the Cjuartzite's. The* teiurmaline is a sliglitly chromi-
fe'rous pale breiwnish schorl with e ~ 1 • (559 + -002, o> = 1*(515 + *002, con-
taining 0-35 pe'r cent (V. 2 O 3 . This oe-curreau'e seems tei support tlie possi-
bility that the^ chromium was elea*ive'el from the granitie- gneiss intrusions
(or freim its pe'gmatitic de'rivative >8 whie*h are ])artie*ularly nume'rous in the
Gorge' Orea'k loe^ality at Ke'y Farm, whea-e the' gre'e'n niusce)\'ite is also more'
abunelant than elsewlu*re).

(b) The Lower Quartzites.- -Them' are' six distine-t banels in the' lower part
of the series. They are inte'rle'ave'el with sillimanite schist, hornbk'iiele schist
(plagioe-lase amphibolite), anel thicker granite gne*iss banels.

92

Rex T. Prider.

They are mostly coarse grained types (average grain size 3 mm. or more)
and have the same granoblastic structures as noted above, the main differences
to the Upper Quartzite being the almost constant presence of felspar in the
Lower Quartzites and its absence in the Upper Quartzite and the fact that tlie
chrome muscovite in the Lower Quartzites is generally set between the quartz
grains while in the Upper Quartzite it occurs as inclusions in the cpiartz.

Most hand specimens are very coarse sugary rocks with bedding planes
sparsely coated with pale greenish chrome-muscovite. White turbid felspar
is a common constituent, but is never present in amounts exceeding five per
cent. It is usually a microcline, but rare gi*ains of oligoclase and orthoclase
are present. The felspar grains are much smaller than the quartz, reaching
a maximum of about 0 * 5 mm. diameter — ^they are usually somewhat rounded,
slightly turbid inclusions in the quartz, but the larger grains show a tendency
to idiomorphism (figure 4A).

Pig. 4.

A. Pelsj>athic quartzite, showing coarse granoblastic structure
and tendency of larger microclines to assert their crystalline
form.

B. ‘‘Heavy mineral’^ band in lower quartzite.

Minor detrital minerals occur sparsely in these lower quartzites — the most
common species are rounded grains of zircon, magnetite and rutile. Pyrite
in rounded grains with a thick rim of limonite is found occasionally— there
can be little doubt, in view of the rounding of the grains, that the pyrite is
an autochthonous constituent of the rock. The segregation of these ‘‘ heavy ”
minerals into bands (figure 4B) has been previously noted.

(c) The Origin of the Quartzites . — The (piartzites, then, are rocks of simple
composition consisting almost entirely of quartz (95 per cent + ). There can
be little doubt that they were originally remarkably pure sands which have been
completely recrystallised in the sillimanite zone. The coarse granoblastic
structures, which have completely obliterated any sign of original clastic
structures, testify to the high grade of metamorphism to wliich tliey have been
subjected. In the absence of any foreign material, the only change in these
rocks is the complete recrystallisation of the quartz. Hall and du Toit (1923.
p. 77) in describing the very coarse quartzites at the base of the Bushveld

Oeology and Pp:trolo(jy of Part of Toodyay District, W.A. 93

Fabric diagrams of tlic .Jimpordiiig quartzites (botli rocks cut
normal to the b-luieations; .

A. Showing rare type of ‘^Trener orientation’’ (Fairbairn,
1937, p. 70) — a marked elongated maximum coinciding with the
“c” fabric axis. Dotted line is plane of foliation. Section per-
pendicular to marked lineation on bedding plane, (.’oiitours 7, 5,
4, 3, 2, 1.

B. Sliowing normal ^^ac” girdle with two prominent maxima,
equally disposed to the ^‘ab” plane. Dotted line is plane of
foliation. Section normal to dij). No lineation a])parent in
liandspecinien. Contours 7, 0, 5, 4, 3, 2, 1.

94

Rex T. Prider.

Complex near Pretoria, are of the opinion that the original purity of the sedi-
ment IS essential for the formation of ^-ery coarse grained types of quartzites.
The purity of the very coarse quartzites described above supports this theory.

That these rocks were deformed in an almost plastic state is seen in the
minor drag folding which they exhibit ^'ocks from these dragfolded areas
show no more cataclasis than in the less disturbed parts.

It IS impossible to recognise any chronological succession in the quartzites
as practically all sedimentary features have been obscured by recrystallisation.

(d) The Fabric of the quartzites and its relation to the geological structure . —
Fabric analyses have been made of several qiiartzites, but in the absence of a
set of geographically oriented specimens, no conclusions can be drawn from
these analyses. They are quoted here to show that the quartzites have a
well-marked girdle fabric (he., are S-tectonites) and also because one of these
rocks (figure 5A) exhibits a rare type of orientation (type “ b ” of quartz
orientation, hairbairn, 193/, p. 70) — the optic axes are concentrated in a
direction normal to the bedding planes {%.€., a prominent maximum lies in
the direction of the “ c ” fabric axis). This maximum is somewhat elongated
and spreads out into the typical girdle.

The other analysis (figure 5B) is a more normal type. In it there is a
well developed ac ” girdle with two prominent maxima, equally arranged
on both sides of the “ ab ” plane. This is a common type of fabric noted
by Phillips (1937) in the Moine Schists of Scotland.

The analyses made, although insufficient to warrant any conclusions,
serve to indicate that a fabric study of these rocks would probably yield much
infoimation regarding the tectonics of this region. One feature brought out
is that the b fabric axis in the tw'o analysed specimens coincides with the
lineations on the bedding surfaces (see figure 2). Although no further fabric
analyses have been made, as no universal stage ecpiipment is available in
Western Australia (the analyses of the two unoriented specimens described
above -were made in the laboratory of the Department of Mineralogy and
Petrology at Cambridge under the direction of Dr. F. 0. Phillips), a re-examina-
tion has been made of certain parts of the area with a view' to determining the
direction and jjitch of these b ” lineations which, on the evidence of the
two fabric analyses described above, are coincident with the “ b ” fabric
axis (ne., the tectonic “ strike ” or tectonic axis, sec Phillips, 1937, p. 587).
These readings are indicated in figure 1, and show that there has been
considerable cross folding on the main N.W. trending structure on axes
trending N.N.E.

Although this structural study of the Jimperding quartzites is far from
complete, sufficient has been done to indicate the desirability of continuing
the work when universal stage equipment is available.

(2) The Calc-Silicate Rocks.

A rather interesting group of lime silicate rocks occurs in narrow kuiticular
seams at a constant horizon in the lower quartzites. The rocks lie just below
the hornblende schist band and are of rather sporadic occurrence. They are
characterised by the presence of grossularite, diopside, pale green amphibole,
epidote, and sphene.

The first sign of the presence in the quartzite of lime-magnesia impurities
is seen in the development of a pale greenish actinolite along with a little
epidote in some felspathic quartzites (e.g., specimen 1250*). Such rocks are

* Numbers are the catalogue numbers in the collection of the Department of Geology, University

of Western Australia.

Geology and Petkology of Part of Toodyay District, W.A. 95

coarse granoblastic structured types in which the amphibole occurs in pale
greenish prisms with irregular terminations, either included in the quartz or
set between cjuartz grains. It all has the same orientation and thus tends to
give a banded appearance to the rock. Rarely, grossularite is noticed as
a constituent of the felspathic quartzites (specimen 1248). Irregular shaped
epidote granules and small lozenge shaped idioblasts of sphene are also in-
cluded in the quartz. In these rocks the (Ca, Mg) silicates are only present as
accessories.

A. Quartz-diopside-epidote-grossularite rock (1249), showing
grossularite aggregates, with associated sphene, epidote and
diopside. The clear areas are quartz xenoblasts.

B. Hornblende-diopside quartzite (15405) witli grossularite
(at top) and carbonate (centre and bottom).

An increase in the dolomitic impiirity in the original sediment leads to the
<levelopment of several types of rock ; — -

(a) Q.uartz-epidote-diopside-grossular rocks.

(b) Hornblende-diopside quartzites.

The former type {e.g., specimen 1249) is a fine-grained, massive, pale yellow^-
ish, homfelsic rock with a banded structure, due to the presence of parallel
seams and lenses of vitreous ciuartz, one or two mm. in length. Under the
microscope the constituent minerals are : — grossularite, epidote, diopside,
quartz, and sphene.

The grossularite is the most abundant constituent forming 00 per cent
or more of the rock. It occurs in aggregates of small idioblastic crystals of
])ale yellow colour. The irregular interspaces are occupied by clear unstrained
quartz, towards which the garnet is always idioblastic, also drop-like quartz
imdusions are common in the garnet. The garnet is all isotropic and has
N 1*70.5 (by immersion in oils) and is, therefore, a grossularite with slight
admixture only of the other garnet molecules.

A pale yellow coloured epidote with very weak pleochroism is fairly
abundant, generally in skeletal crystals which, except for the development

96

Rex T. P rider.

of a good 001 eleavag(‘ and poorer 100 cleavage, are almo.st indistinguislmhle
from the grossularite under ordinary light. The refractive index /3 clos(‘ to
1 • 765, extinction Z /\ 001 cleavage 30°, y-o approximately -042 and neg-
ative optical character, indicate an epidote with approximately 15 per cent
(Winchell. 1933, p. 313). TIichc skc'letal crystals often enclose idioblastic
grossularite.

Diopside is not so abundant. It is a pale greenish variety in hand sp(‘ci-
men, colourless in thin section. It occurs in a similar fashion to epidote,
enclosing grossularite. The greater part of the diopside is confined to the
quartzose portions of the rock, where it is idioblastic towards the (piartz.

Sphene is an abundant accessory in cloudy irr(‘gularly shaped allotrio-
morphs usually surrounded by gariif't. Accessory opacjue iron or(‘ occurs
in several bands.

Quartz, the only other constituent, occurs in elongat(‘d xenoblasts and
skeletal grains between the garnet ciystals. There are no signs of any sti-ain
in the (piartz.

The crystalloblastic order is : spliene, grossularite, (‘pidote, diojxside,

and (juartz. The microstructures are shown in figure 6A. An analysis of this
rock, quoted from Sim})son (1937, p. 32) is

SiO., ...
TiOa

AhO.

FeaOs

FeO ...

MnO

MgO

CaO ...

Alkalies

H.O+

H2O-

50 • 68

0 - 96
12-63

3-79

2-13

1 - 02
1 -71

26-81

Tr.

0-35

0-26

100-34

Judging from its high lime and alumina and absence of alkalies the j-ock
was originally an impun* siliceous and argillaceous limestone. The C’aO,
AI2O3, and SiO., liave given grossularite, the entrance of some Fe20;3 has letl
to the developna^nt of epidote and the C'aO with the small amount of MgO
has given rise to diopside. The source of the titanium of the analysis is spheiu',
of which there is a small amount in the rock.

The hornblende-diopside quai'tzites are represented by a numb('r of
.specimens in the collection of which 15405 shows all the charact(a‘istic feature's.
Megasco])ically, it is a coarse-grained, dark green (juartzite. Under the micro-
scope, it is (‘ssentially a coarse granoblastic aggn'gate of ejuartz grains (2 3
mm. diameter) which carry poikiloblastic inclusions of pale greenish diopside^
(Z/\c 42° indicating the presence of some iron) and greenish amphibole,

Aggi‘ogat('S of diopside, hornblende, grossularite, and a little carbonate occur
between the quartz grains. The (piartz (70 per cent of the rock) is xeno-
blastic towards all other constituents. It is all considerably strained and
carries minute gas-li(piid inclusions.

The amphibole, occurring in pale greenish .sheaf-like aggregates, ajip('ars
to be developing from the diopside which is occasionally seen as relicts in the
amphibole prisms, d'he chai’acters of this amphibole are : —

Pleochroi.sm X pale yellow-green ; Y deep olive green ; Z bluish-
green.

The absori>tion is Y>Z>X and Z/\c = 19°, indicating that it
is blue-gret'ii hoj'nblende rather than actinolite.

Geology and Petrology of Part of Toodyay District, W.A. 97

Grossularite is present in small amount only, in irregular shaped, isotropic,
yellow granules, always closely associated with the diopside. There is no
epidote in 15405, but it occurs in similar rocks and possesses similar characters
o that in specimen 1249 above.

15405 is the only rock in which any carbonate has been noted. It occurs
in small xenoblastic forms, always with concave boundaries towards the
other constituents. It is most abundantly developed in the vicinity of diopside
and hornblende (figure 6B). The complete absence of twinning suggests that
it is dolomite rather than calcite.

There can be no doubt that 15405 and related rocks are meta-dolomitic
sandstones. Failure of the carbonate to react completely with the silica
appears to indicate lack of sufficient aluminous and ferruginous impurities in
the original sediment. The co-existence of quartz, grossularite, and calcite
(or dolomite) indicates that these rocks have been developed by regional
metamorphism under high pressure, for, if thermal metamorphism were
responsible, then it would be expected that the remaining carbonate would
liave reacted with silica to give wollastonite.

The rocks described above are representative of the sparsely distributed
calcareous seams in the original sandstones. They undoubtedly represent
T'('gionally metamorphosed impure dolomitic sandstones with a varying car-
bonate content. Kaolinic and ferruginous impurities have led to the forma-
tion of epidote and grossularite and a certain amount of iron has entered
into the dolomite-quartz reaction producing diopside. In rare instances an
insufficiency of impurities has inhibited the dolomite-quartz reaction. The
grossularite in these rocks appears to have developed under conditions of
regional metamorjDhism in the sillimanite zone. Tilley (1937, p. 372) has
<lescribed the development of grossularite under stress conditions in the Loch
Tay limestone of Perthshire. Here the grossularite, together with vesuvianite
and diopside, first makes its appearance in the Almandine Zone. The Tood-
yay grossularite rocks show^ remnants of epidote, which in the Loch Tay
limestones appears earlier than grossularite and disappears in the deeper
})arts of the Almandine Zone. The presence of epidote in the sillimanite
zone at Toodyay, along with grossularite, indicates that epidote may, under
<-ertain conditions persist through the Almandine Zone.

(3) The Mica ScTists.

These politic members of the Jimperding Series occur at two horizons.
In the upper part of the series, a band of mica schist, approximately 250 feet
thick, forms the upper limit of the series. This band is characterised by
al>undant andalusite. Another thinner band near the bottom of the series
is characterised by sillimanite. Throughout the whole area the mica schists
m-(' considerably weathered, making examination o fthem somewliat difficult.

The lower jnica schist can be ti’aced from the south-east corner of the
ai-ea alniost to the western side. It is interbedded witli (piartzites and is
<‘\'erywhere c.ontorted by small dragfolds consequent upon its having been
t()ld(^l between the two more competent quartzite beds. In the central
part of tlie area and in the immediate vicinity of the townsite of Toodyay,
it is considerably thickened by folding and it has l)oen impossible in this
<'rumpled portion to map all the minor folds.

98

Rex T. Prider.

The following types are developed amongst the mica schists : —

(a) Sillimrmite Schists . — These are mainly two-mica schists with narrow
psammitic bands. Slight variations have been noted and they are illustrated
by the following descriptions of typical members :

(i) Quartz-biotite-sillimanite schists (e.g., spec. No. 15638). The‘s<^
are finely schistose structured rocks consisting of bands composed
almost entirely of fine acicular sillimanite, alternating with quartz-
biotite bands which represent more ferruginous and psammitic
layers in the original sediment. The fibrolite bands are occasion-
ally distorted by a later growth of sillimanite in stout cross fractured
prisms arranged at about 45° to the schistosity (figure 7A), in-
dicating that a later thermal metamorphism was imposed on the
earlier regional metamorphism which had developed the fibrolite.
Muscovite (rare) also has a tendency to be arrang(‘d with its
cleavages at about 45° to the schistosity.

Fig. 7.

A. Sillimanite schist — bands of fibrolite with a little biotite,
alternating with quartz-biotite bands. A later growth of silli-
manite developed at about 45® to the banding. Note ])sammitic
quartz-biotite bands.

B. Sillimanite-biotite-plagioclase granulite. No tendency to
orientation of any constituents. The clear areas arc fine granu-
lar plagioclase.

The fibrolite appears to be developing from biotite, the iron released
in this change being represented by elongated grains of magnetit('
which occur most abundantly in the fibrolite layei’s.

(ii) Sillimanite-biotite-plagioclase granulite (c.g., spec. 15078). Such
I'ocks as these show’ the very interesting association of sillimanite
with zoned plagioclase and myrmekite. In hand specimen the
rock is tine-grained, greyish in colour ’with an irregular banded
structure — fissile biotite bands alternating with granular quartz
fels])ar bands. Under the microscope the structure is fine, even-
gi’ained granoblastic w’ith lenticular areas rich in sillimanite and
biotite with schistose structure. The approximate mineralogicaj

(]1eology and Petrology of Part of Toodyay District, W.A. 99

composition is quartz 10 per cent, oligoclase 35 per cent, silli-
manite 20 per cent, biotite 30 per cent, muscovite 5 per cent,
and accessory magnetite and zircon, but in certain bands the
sillimanite content rises to 70 per cent or more of the rock which
thus becomes a valuable sillimanite ore.

In the psammitic bands both c(uartz and oligoclase are xenoblastic.
I’he felspar is invariably zoned, the zoning being reverse gradational
in character from Abg An^ centre to Ab 3 An^^ periphery ; twinning
is seen in the central parts but the outer zone is generally un-
twinned. The outer zones of the plagioclase grains also carry
numerous vermicular inclusions of (juartz, forming a myrmekite-
like structure. The development of this structure in the absence
of microcline is interesting — it appears to be due to the growth
of the plagioclase around crushed (juartz spindles.

The sillimanite is in two generations (1) as fine acicular clusters,
(2) stout cross fractured rods, the arrangfunont of which is unusual
(figure 7B). In any section they appear to be arranged in bundles,
some' lying in the plane of the slice, others arrangcnl normal to or
at an angle with it.

The biotite is a deep rc^ddish-brown lepidomelane in im‘gular shaped
flakes with but little tendemey to parallel orientation. In addition
to inclusions of sillimanite there are numerous minute zircon
inclusions, surrounded by pleochroic haloes.

Muscovite is in larger plates (Imm.) carrying poikiloblastic in-
clusions of (juartz and oligoclase. It is pc'netrah'd by sillimanite
rods but is idioblastic towards the biotite.

Magnetite and zircon are the only accessories.

(iii) Cordierite-sillimanite schists {e.g., specimen 15683). This sj)('cimen
is the only metasediment of this area in which the preseiKfe of
cordion'ite has been noted. Ther(‘ is no sign of schistosity in the
rock, either in hand specimen or in the field occuirix'nce. Con-
stituents noted microscopically were quartz, biotite, muscovite,
sillimanite', and pinite (after cordierite). The sillimanite is found
as small tufts of needles closely associated with a deep rc'ddish-
lirown biotite and radiating out into the neighbouring ejuartz.
This variety is abundant and appears to be developing at the
('x])('nse of the biotite. Sillimanite occurs in a second g('neration,
in stout('r rods haphazardly includc'd in a pale brownish practically
isotropic pinite (after cordierite) ; in section these jiinitic areas
mc^asure up to 0-5 mm. diameter. There is no definite oric'nta-
tion in the micas. Plagioclase is absent.

The ajiproximate mineralogical composition is -biotite 35 p('r cent,
sillimanite 25 p('r cent, (juartz 25 p(T ccnit, pinite U) jier ceiit,
muscovite five p(T (^ent, and ac(^('SSory zircon and magnetite.

(6) (kirnvt-hiotite-plagloclase schist.- occunence of garnet in the mica
schists has only been noted in one place (15 cliains >S., 122 chains \V. from
datum). H('r«i the garnets are found in a very sandy schistose rock, which,
under the microscope, is seen to consist largely of zoned plagioclase. The rock
is medium grain('d, schistose and granular in structure, with porphyroblasts of
pink garnet up to 3 mm. diameter. The greater part of the rock consists of an
ecjuigranular completely recrystallised aggi*egate of jilagioclase, cjuartz, and
biotite with an imperfect schistose structure due to tJie subparallel alignment

100

Rex T. Prider.

of biotites. The scliistosity is interrupted by the idioblastic garnets which,
during their growth, have pushed the biotite flakes aside. Tlie crystalloblastic
order is garnet, biotite, plagioclase, and quartz.

The biotite is a strongly pleochroic variety with X pale yellow, Y = Z
<.lecp brown, and refractive indices, y = 1-G38 and a = 1*591. Zircon in-
clusions whth pleochroic haloes are abundant. The plagioclase is an albite
about AbgAn^ and shows slight normal gradational zoning. The centres of
some of the crystals are sliglitly turbid wutli sericite-epidote alteration products.

-r 8 Feet

o O o

o o o o

O O o

O O

O O O

o o

d 0 - ^ e

o 5

O o

Mica schist
Quartz-mica schist

Garnet-mica schist

Cummingtonite schist

Garnet-mica schist with
cummingtonite hands (§” -wide)

Mica Schist

Fig. 8.

Sketch section of small w^aterfall at the head of Gorge Creek
(15 chains S., 122 chains W., from datum), showdng the relation
of cummingtonite schists to the mica schists.

From the same outcrop as tlie garnet schist comes : —

(c) Gedrite-cummingtonite-plagioclase schist . — A typical specimen (19947)
is a pale greenish-grey, fine grained bandcnl rock- — light greyisli to white' bands
to J in. tliick alternating witli gr('enish-grey bands up to \ in. tliick. Under
tlie microscope there is a well-marked schistose structure and the minerals noteil
w'orc gedrite, cummingtonite, plagioclase, and c[uartz.

The amplhboles occur in well-shaped prisms (often w'itli ])oikiloblastie
inclusions of plagioclase) uj) to 1 • 5 mm. in length. Tliey all he witli their
longer axes in the plane of scliistosity, but appear to be of random arrangement
in these planes since basal and longitudinal sections are present in aj^proxi-
matciy eipial numbers in all sections cut at right angles to the scliistosity.
Both are jiale clove-brown, not noticeably pleochroic varieties and are dis-

(Ieology and Petrology of Part of Toodyay District, W.A. 101

t ingui.shable only by the inclined extinction (Z/\c = 17°) of the oiimmingtonito
as (compared with the straight extinction (Z = c) of the gedrite. The former
oft(‘ji shows lamellar twinning. The optics of the latter : —

Pale clove-brown, non-pleochroic, X = a, Y = b, Z -- c, (-) 2V
large, y — DOOV, ^8 = 1*659, a = 1*653, indicate the aluminous
variety, gedrite.

'Fhe light-coloured bands are composed of cummingtonite (with gedrite)
- plagioclase -|- ({uartz, while the darker-coloured bands are much richer in
cummingtonite and contain little or no ({uartz, and in some still darker bands
gre(‘n hornblende becomes an important constituent.

vS])ecimens transitional in character between (6) and (c) are present in the
same outcrop (see figim^ 8), and in these specimens narrow dark bands of
garnet-biotite-plagioclase schist (often containing prisms of g('drite) alternate
with light-coloured cummingtonite plagioclase^ rock.

A noti(teable feature is that the ampliibole in the biotitic bands appears
to b(‘ all gedrite, whereas this mineral is subordinate to cummingtonite in
the lighter-coloured layers.

All the above types («)» (^)» (c) found in the lower band of mica

schist. They have been derived from argillaceous sediments of variable
composition ; plagioclase becomes evident in the slightly calcareous bands.

The various associations of sillimanite, biotite, (juartz, plagioclas(‘, and
cordierite noted above are all well known. The g(^drite-cummingtonite-
biotite-oIigoclas(^ association is much rarcT. Amongst the best known com-
parable rocks are -

(!) The gedrito-plagioclasites of the Kragero region, Norway, described
by Brogger (1935, ])p. 213-325). Although generally massiv(',
schistose types have been described from this district. Thcvse
rocks ar(' (jonsidered to be derived from basic igneous ro(*ks in a
similar fashion to the gedrite (and anthophyllite)-cordiorite rocks.

(2) 'Phe ])lagioclase-gedrite gneisses of the Nesodden Peninsula n(‘ar Oslo,

described by Brock (1926, p. 180) in which the association “ (juartz,
plagioclase, biotit(', garnet, gedrite agrees very closely with that
(>bserv(*d in the Toodyay rock. Brock considers that these rocks
have b(‘cn derived from leptites by addition of FeO and MgO.

(3) 'Phe garnet plagioclase amphibolites of Isopiia, Kalvola, Finland,

(l(‘S(Tibed by Fskola (1936, p. 475). P^skola in this pa]K‘r d('scribes
a paragenesis of gedi’ite and antho])hyllite, an association similai*
to that noted in the Toodyay schist, and he considers that the
g(‘drite-cinnmingtonite-garnet amphibolite of Isopaii has becui
formed by metasomatic replacement of basaltic agglomcTates.

'Pile gedrite-cummingtonitc’-biotite-plagioclase schist from Toodyay, from
its int('rlamination with sillimanite schists and tlu' rhythmics banding obs(‘rved
in the specimens appears to be a metascHlimentary rock, deri\’('d probably
from a tliin bedded sediment such as an ini]')ure dolomitic shal(‘.

{(1) Andahjsite schists. -These are confined to the uppermost })elitic band
which outcrops in the S.W. portion of the ar(‘a. The rocks of this band are
(juartz-muscovdte schists with a variable biotit(‘ cont(ait. All the outcro])s
are highly weatla'red, the rocks aj)pearing as yellow and reddish iron-stained
schists. Simpson (1936, p. 11) has noted the occurrence of sillimanite (fibro-
lite) in these andalusite schists but it appears to be very rare, as I have noted
the presence of sillimanite in one specimen only.

102

Rex T. Prider.

The aiidaliisites sliow a constant variation in size and number tliroughout
tlie band as shown in figure 9. At the bottom of the band the andalusites
are very abundant — they project from the weathered surfaces forming 40 per
cent or more of the exposed surface. The crystals vary in size up to 6 x 2 x
2 cms. The prism (110) is always well developed but the terminal faces are
very poorly developed.

Traversing the band towards the south the andalusites become much
smaller and less abiindant and finally disappear, the rock then being a much
weathered reddish muscovite schist which outcrops right up to the granite
boundary.

K

A^/? 7Z//? L ScfIL £

1 1

1000 F'ecf

Pig. 9.

Cross section of upi:)cr mica schists showing distribution of nndnlusitc.

The andalusites in the lower part of the band, as described and figured
l)v Simpson (1928, p. 50), all show some alteration to muscovite : —

(1) On the periphery, to a coat of small muscovite plates.

(2) Within the crystals there is usually an irregulai- alteration to fine

fibrous sericite. Most of the andalusites are clouded with carbon
dust inclusions and show the typical chiastolite cross. In addi-
tion, we Tiiay note' a peculiar feature shown by many of these
andalusites : the arms and cleavages of the chiastolite cross are

curved but tiie optic ori^nitation remains constant throughout the
wliole crystal. This feature is similar to that shown by some
staurolite and garnet crystals (Harker, 1932, p. 221) due to rota-
tion during their growth.

Tlie groundmass of these rocks is a ijuartz-biotite-muscovite schist. The
biotite is completely wt'athered. It is interleaved with coarse platy musco-
\-ite. Quartz is abundant in sTualler irregular shaped grains. Plongated
lenticles of sericite are common. Tliis material is similar to tliat developing
in tlie larger andalusites and it appears to result from the shearing out of
smaller andalusites in the bas(' of the rock.

In one specimen this sericitic^ matei’ial is present in lenticular areas with
iri-egular inclusions of quartz. The micas are deflected arountl these knots in
a similar manner to the deflei'titm around larger andalusite porphyroblasts.

Geology and Petrology of Part of 'J'oodyay District, W.A. 103

Tlie only other constituent is rare, w^elhshaped brow'iiish tourmaline.
3n one specimen a few stout rods of sillimanite, arranged parallel to the folia-
tion were noted, but this is the only sillimanite present in my specimens.

Proceetling in a southerly dirc'ction towards the granite, the andalusites
become smaller and eventually disappear. Specimens from this part are
similar to the groundmass of the andalusite schist but have a greater abundance
of the lino fibrous sericitic bands and knots, lirownisli tourmaline is present,
l)ut rare. This type of rock persists right up to the granite contact which
cuts the schists off abruptly to the south. In a series of specimens collected
throughout this band there was no apparent variation in the tourmaline
content. The absence of any concentration of this constituent in the schists
near the contact seems to negative the possibility that it was introduced from
the granite. At the same time its constancy of character indicates a common
source for all the tourmaline. Its origin is therefore somewhat doubtful, as
it may be developed from constituents introduced from the intrusive granite,
07' it may be original detrital toui-maline wliich has been recrystallised.

In considering the origin of the andalusite we naturally look first to the
intrusive granite as the source of heat. The granite is intrusive because : —

(1) It ti*ansgresses the bedding of the mica schists.

(2) Several patches of schists up to 20 cliains long x 8 chains wide

liave been noted wliich are surrounded by granite.

(3) It shows no foliation like the granitic rocks in the Jimperding

S('ri(‘s.

Tliere has been no apparent thermal alteration of the (country I’ocks.
As shown above, the andalusite^ is abs('nt in the vicinity of the contact and,
therefore, the genesis of the andalusite could only be attributed to thermal
alteration by the granite if there were an increase in the pcilitic constituents
of th(? original st'diment as it becomes more distant from the granite. Althougli
no chemical analyses are available, microscopical examination shows that
tlie composition is faiily constant throughout the band, and this rules out the
granite as the agent which formed the andalusite.

The granite' on its outer mai’gin has sufferc'd no chilling and is just as
<*oai'se-grain(‘d as it is at a distances of a (juarter or half-mile from the contact.
'^Phis suggc'sts that the intrusion of the granite took place at some depth, into
se'diments which were themselves in a somewhat heated condition. The
])ossibility of a faulted junction between the granite and schist must, however,
not l>e ov'erlooked.

The distribution of the andalusites indicates an accession of heat from
lu'low. There are sevei-al inti’usive masses, below the andalusite' scliist (figure
9), which may have su])pli(‘d the lu'at lU'cessary foi' the dc'velopment of andal-
usite :• •

(1) Th(* Uppc'r Gi'anitic giu'iss.

(2) An ultrabasic sill now rejresentc'd by a monomineralic actinolite

schist.

(3) Doleritc dykes. Theses could not have* (‘ffectc'd tlu> devoloj)m('nt

of andalusite as they ai’c post granite in age. wlic'i’c'as the develop-
ment of andalusite was pre-gi’anite.

Latc'r retrogi'c'ssive clianges in the andalusite (i.c., replacc'ment by musco-
vite and sericit(') are ])robably due to pneumatolytic altc'i’ation by va])oui's
from th(‘ intrusive granite to the south, in a similar manne]’ to the alteration
of sillimanite'- to muscovite noted by Simpson at Clackline (Simpson, 193().

104

Rex T. Prider.

p. 13). In addition there has been a considerable amount of sheai-ing aft(a-
the formation of andalusite, which in specimen 10743 is cracked and slieared
out into lenticles consisting mainly of sericite but carrying relicts of andalusite.

(4) Hornblende schists (plagioclase amphibolites).

There are several different modes of occurrence of these rocks ; —

(1) as well defined bands (average thickness 40 feet), interbedded

with quartzites. All have been completely recrystallised and
variable grain features, which may indicate the original nature
of the mass (Cooke, etc., 1931, p. 49) have been obscured. These
bands never transgress the bedding of the associated sediments,,
and they appear, in view of the great area ovt'r wliich they
retain their constancy of character and liorizon, to l)e basaltic
flows which have been folded along with the associated sediments.

(2) as irregular shaped inclusions, usually of small areal extent, in the

upper granitic gneiss. These are considered to be xenolithie
bodies.

(3) coarser-grained xenoliths in tlie lower granitic gneiss. These are

often veined with granitic material.

The two latter occurrences will be considered later in a section dealing
with the xenoliths in the gneiss.

The schistose plagioclase amifliibolites interl)edded witli the metasedi-
mentary rocks are remarkably constant in character, consisting mainly of
blue-green hornblende and ac'id plagioclase, with minor amounts of e})idote,
(juartz, microcline, magnetite, s])hene and apatite. The only variants of
this type are diopside-plagioclase amphibolites of rather rare occurrence.

15437 is a typical example of the normal liornblende schist. It is a
medium grained schistose rock consisting predominantly of hornblende.
Epidote is an abundant constituent, and in hand specimen is seen to occur
in pale yellow-green seams running ])arallel to the schistosity.

Under the microscope the rock has a well defined schistose structure,
the hornblende occurring in irregular shaped prisms (average 0-5-mm. long),
in parallel alignment. Other constituents are plagioclase, microcline, epidote,
(piartz, and sphene.

The hornblende j^risms often carry small ])oikiloblastic inclusions of
([uartz and felspar. The liornblende is a common blue-green variety with
X yellow to yellow-green ; Y olive to brownish-green ; Z bluisli-green, and
absorption X Z Y, ^ ^ 1-072 and Z/\c = 20°.

The epidote is the pale yellow highly birefringent pistachite, usually
confined to narrow bands parallel to the foliation. In the vicinity of epidote
the hornblende is represented by a pale greenish bleached variety. Along
joint cracks which are not coated with e]jidote tlie liornblende is also bleached
— such joints traverse the hornblende crystals without dislocating them and are
only evidenced by the jiresence of a narrow seam of more fibrous bleached
hornblende which has the same optical orientation as the crystal traversed.
There has clearly been some transport of material along these microsco[iical
fractures, as seen by the occurrence of the same jiale amphibole along the
joint plane . where it traverses ([uartz and felspar grains.

Two varieties of felspar are present, both xenoblastic an<l slightly
elongated parallel to tlie schistosity. Albite, which is slightly turbid with
fine granular epidotic and fibrous sericitic alteration, is most abundant. It
rarely shows lamellar twinning, is not zoned, and its refractive index y ^ 1-54
indicates an albite with less than 10 per cent An.

Geology and Petrology of Part of Toodyay District, W.A. 105

The less abundant microcline is all water-clear and shows the character-
istic cross hatched twinning. Microcline although rarely present in amounts
exceeding 5 per cent is a constant constituent of these rocks.

The only accessory is sphene, in small imperfect crystals usually en-
closed by hornblende.

This rock has been analysed as typical of these hornblende schists (Table
1, col. 1). It is an analysis closely resembling that of igneous rocks of the
composition of quartz dolerite. An analysis of a dolerite from this area
appears in column 4.

TABLE 1.

Analyses of Basic Rocks from fhe Toodyay Area.

SiOg

A1,03

Fe^Os

FeO

MgO

CaO

Na^O

K^O

H 3 O+

H.,0-

TiO,

P2O;

MnO

b^eS.>

Norms.

Q

Or

Ab

An

di

iiy

mg

il

ap

py

1 .

2 .

3.

4.

53-61

50-20

49-05

49-13

10-77

15-00

15-03

13-13

2-21

3-83

3-16

3-65

9-02

8-93

9-08

8-95

6-92

6-04

6-96

7-64

10-04

10-65

10-47

11-84

2-12

1-90

1-70

1-72

1-54

0-07

0-95

0-16

1-62

1-62

1-43

1-72

0-17

0-07

0-25

0-04

1-05

1-06

1-07

1-27

0-17

0-12

0-08

0-14

0-35

0-16

0-09

0-15

n.d.

n.d.

0-13

0-45

99-59

99-65

99-45

99-99

5-10

6-18

1-98

3-48

8-90

0-56

5-56

1-11

17-82

16-24

14-15

14-15

15-57

31-97

30-58

27-52

27-30

16-41

16-68

24-88

17-52

19-18

21-58

18-33

3-25

5-57

4-64

5-34

1-98

2-13

2-13

2-43

0-34

0-34

0-34

0-34

—

—

0-13

0-45

1. Schistose plagioclase amphibolite (15437), interbedded with quartzite, Toodyay, W.A.

2. Schistose plagioclase amphibolite (1241), xenolith in Upper granite gneiss, Toodyay,

W^. A.

3. Coarse granular quartz-plagioclase amphibolite (15441), xenolith in Lower granite

gneiss, Toodyay, ^^^A.

4. Quartz dolerite (15424), dyke in metamorphic rocks, Toodyay, W.A.

All analyses by R. T. Prider.

Amongst the rocks forming this band there is, as j)reviously noted, but
little variation. Microcline may be absent and ([uartz may be more abundant
than in the tyj>e rock. Iron ore rimmed with spliene, indicating its origin
from ilmenite, is a common feature, although in the rock described above
the usual (‘entral ore inclusion in the S])hene is absent.

T'he most unusual variant of the plagioclase amphibolites is a type in whicli
diopside finds considerable development. 15440 is typical— it appears in
hand sjjecimen to be a normal hornblende schist with narrow bands and
lenticles (never more than 2 mm. wide) of light greenish diopside.

Kkx T. Pridkr.

I OH

Under tlie microscope the constituents are hornblende, diopside, and
jdagioclase with accessory (juartz, apatite, and sphene. The hornblende
and diopside are concentrated into alternate bands (figure lOB) and appear
to have crystallised independently. The hornblende has a much stronger
absorption than that described above — it has X yellow-green ; ’N' brownish-
green ; Z deep bluish-gi’een : and absorption X Z ^ Y ; ^ ^ ^ I ■(>74 ; and
Z/\c — IH°. The diopside is in irregular cf|ui(limensional grains to 0-5 mm.
diameter. They have no <lefinite orientation, tlie longitudinal cleavage being
arranged at various angles to the schistosity, indeed often normal to it.
This mineral is a pale greenish variety with the (110) cleavage well develof^ed,
Z/\c — 42°, indicating the considerable iron content. The dioi)side is-
idioblastic towards both the plagioclase and hornblende. F^lagioclase forms.

Fig. 10.

Pyroxene-plagioclase amphibolites.

A. Coarse xeiiolith in Lower Gneiss. Constituents: diopside
(dotted), hornblende (]>alor coloured at junction with diopside),
slightly turbid plagiocdase, quartz.

B. Fine banded variety interbedded with (]uartzite. Constituents:
hornblende in well defined bands, diopside (dotted), plagioclase.

approximately 30 per cent — it appears in two generations (1) a very turbid
indeterminable variety clouded with secondary fine granular zoisite — this
variety is more or less confined to the hornblendic bands ; (2) a clear fresh
]‘)lagioclase associated with the diopside bands. Fine lamellar twinning
is occasionally develo])cd and the small extinction (2° - 3°) in sections X 010
indicates an oligoclase. Slight normal gradational zoning is notic'cable in
some grains.

Sphene and rare apatite are the only accessories. Iron ores are absent.

Another specimen belonging to this group has an *' augen ” structure —
the augen are of diopside aggregates set in a plagioclase amphibolite ground
in which the felspar is heavily saussuritised.

A specimen taken from the iipper contact of the hornblende schist with
the quartzite at a point 132 chains E. and 229 chains S. of datum, shows a

Geology and Petrology of Part of Toodyay District, W.A. 107

gi'adation from normal plagioclase ami^hibolite. tlirougli epidosito to a diop-
side-]>lagiocla.se rock in which grossularite is ])rescnt in small amount- — this
latter type is developed right at the contact.

The j)lagioclase amphibolite portion of the specimen is the normal type
in which the felspar is very turbid with fine granular e])idote. Following
this is a narrow band (} incli) consisting [predominantly of yellow epidote
(pista<‘hite) with a minor amount of pale greenish hornblende and quartz —
the hornblende then decreases in amount leaving an almost ])ure epidote
band (] incli) in which lozc^nge shaped sphenes are a notable constituent.
In tlie outer part of this band grossularite is occasionally seen. Jkwond
this, the rock consists of epidote, slightly turbid plagioclase (Abc)Anj) and
(juartz, tlie two latter occurring in micrographic intergrowth. On the outer
limit of the specimen, the rock consists of diopside and saussuritised plagio-
clase with a little grossularite, sphene, and ([uartz. Were it not so clearly
related to the [plagioclase amphibolite, siu'h a rock could conceivably have
resulted from the metamor[)hism of an im[pure argillaceous limestone. The
develo[)ment of grossularite in this rock is interesting, as it lias developed
from the epidotic alteration of the original felspar.

The main features which elucidate the origin of these schistose [plagioclase
nmplubolites are : —

(1) They occur interbedded with metasediments and never transgress

the bedding.

(2) They are constant in character throughout the area. Rarely,

more calcic diopside layers alternate regularly with layers of
the normal amphibolite. A similar feature to this is seen in the
Landewednack hornblende schists of tlie Lizard, Cornwall
(Flett and Hill, 1912, [). -M>).

(8) The refractive index — l-(>73) of tl le hornblende is indicative
of hornblendes from e]pidiorites in the sillimanite zone (Wiseman,
1934, [p. 394).

(4) In their chemical com[Position they are normal igneous rocks,
such as would result from the crystallisation of a (] uartz dolerite
magma.

i'lu'se [plagioclase am[Phibolites a|>[pear, therefore, to be m.etabasic igneous
rocks, v'hich formed either a sill or flow in the metasediments [prior to tlie
fprogenic [period, and have been folded along with the metasediments. In
view of the complete recrystallisation’, no evidence is available as to whether
thes(‘ were originally flows or sills.

(5) The (hrmtfe Onetsses and iJte associated xenoliths.

A^. d(‘s(;rib(-d above, tlien^ are two bands of light coloured gmnitic gneiss
<levelo[H'd in the area. The U[Pper and Lower Gneisses form bands estimated
to h(‘ a[)[proximately 2,000 feet and 5,400 feet thick respectively. Minera-
l(pgir-all\-, there is but little difference between tlpe two. although the up[)er
baud, in view of the greater abundance of microcline is somewliat tlie richer
m [Potasli. It has also suffered more crushing than the Lower Gneiss and is
<‘V(Tywhere a ty[ji(*al augen gneiss with large " eyes ” of microcline, sur-
rounded by a granular ( juartz-biotite-microcline aggregate. The lower gneiss
IS essentially a fluxion gneiss, in many [places a coarse [Porphyritie ty[Po with
idiomor|)hic microcline [phenocrysts (iqp to I inch long), which have a linear
orientation due to flowage. Both gneisses are penetrated }py late stagi'
[pegmatitic [products of the granitic magma— these may be narrow sills or
\eiiplfts traversing the foliation, usually only a few inches wide.

108

Rex T. Prider.

The Upper Gneiss is constant in conaposition and structure throughout.
In places the crushing has been so complete as to obliterate the augen
structure and fine banded gneisses have resulted. Such rocks have elongated
narrow lenticles rich in fine granulated microcline, alternating with long
sill-like streaks of unstrained c^uartz, in which the cpiartz extinguishes as a unit.

The foliation in both bands conforms to that in the associated meta-
sediments. In the case of the upper gneiss, the gneiss-quartzite contact is
often a narrow crush zone, varying from a few inches to a few yards in v idth.
The rocks of this zone are (juartz-sericite schists, in which the cpiartz appears
in elongated lenticles up to 1-| inches x } inch x J inch with prominent cross
fractures (tension joints) — these lenticles are elongated in the plane of
foliation, but more elongated parallel to the strike than to the dip.

In both gneisses there is often an irregular folding and contortion of
the flow layers rich in biotite, due to folding movements which were taking
2 >lace contemporaneously with the flowage.

That the gneisses are of igneous origin cannot l>e doubted in the evidence
presented by mimerous xenolithic bodies of variable composition— some
metasediments, others metabasic igneous. These will be described presently.
In places at the bottom of the Upper (hieiss (notably in Poison Creek), the
gneiss is discordant to the bedding in the (piartzite and small offshoots
traverse the bedding of the ({uartzite.

We will consider firstly the Uppei' Gi’anitic Gneiss and its associated
xenoliths.

(a) The Tapper Granitic Gneiss.

The niaiji characteristics may be summarised thus : —

(1) Augen gneissic structure is characteristic throughout.

(2) Ptygmatic folding is occasionally developed.

(3) With the exception of occasional small basic xenoliths, narrow

lenticles of plagioclase amphibolite and irregular small biotite

granulite xenoliths, the composition is fairly constant throughout.

(4) Microcline is the dominant felspar.

(5) Myrmekitic structures are developed in most types.

(6) Biotite is im-ariably chloritised and contains sagenitic rutile in-

clusions.

(7) Narrow veins of pegmatite and aplite are numerous.

(8) Garnet and the ^•a^ious aluminium silicates characteristic of meta-

sediments are completely absent.

The main type of the Upper Gneiss is a microcline granite gneiss, but
other minor types are occasionally noticed. The varieties of Upper Gneiss-
are : —

(i) Biotiteanicrucline granite gneiss. — These are medium-grained rocks
vlth augen gneissic structure. The augen are of microcline up
to 1 cm. X O’O cm., in an even-grained ground of cpiartz, felspar,
and biotite, the latter in sub-parallel alignment. The cpiartz is
in elongated ellipsoidal grains (5 mm. x 3 mm. x 1 mm.) showing
greatest elongation parallel to the strike and lesser parallel to the dip.

Under the microscope the minerals observed were quartz, micro-
cline, oligoclase, and biotite vith accessory magnetite, apatite,
muscovite, zircon, epidote, rutile, and myrmekite.

Microcline is most abundant, occurring in xenoblastic plate‘S, usually,
but not always showing peripheral granulation (Plate III, C). This

Geology and Petrology of Part of Toodyay District, W.A. 109

granular microcline aggregate is comparatively coarse-grained and
seems to be a protoclastic, rather than a cataclastic structure —
this is supported by the absence of any cataclasis or marked strain
in the associated quartz. Tlie microcline may carry small inclusions
of quartz and plagioclase, and in the latter instance mynnekitic
intergrowths of quartz with the plagioclase are usually present.

Pig. 11.

Fabric diagram of Upper granite gneiss. Plot of
poles of optic axes of 200 quartz grains, showing the
complete absence of any orientation in the quartz.

The Cjuartz is in irregular shaped unstrained grains forming a con-
stituent of the microcline mosaic and, as alloti’iomorphs, elongated
parallel to the gneissic banding (Plate IT, A and B, and Plato
III, D), in which case it often includes parallel aligned biotite
flakes. The (juartz shows slight undulatory extinction but has
not suffered any granulation and has been either : —

(1) of post t(*ctonic crystallisation : or

(2) original cruslied gj’ains, eoTnphdely recrystallised into elong-

ated xenoblastic forms.

A fabric analysis was made of a fine even-grained granitic gneiss,
showing this marked elongation of unstrained ijuartz grains, but
after the measurement of 200 grains no apparent concentration
of the o])tic axes in any direction was noticeable (figure 11).
Similai' results wert‘ obtained from se\-eral othei* granite gneiss(>s
wliich wei'(‘ analysed.

It is interesting to coinpaie this result with those obtained for several
of the (juartzites (figure 5. A and B), which show sucli a well-
marked fabric. It would be expected that the gneiss would
possess a fabric similar to the (piartzites. The complete absence
of any girdle in the diagram for the gneiss indicates that the quartz
is of post tectonic crystallisation. This is verified by the observation,
in somf' j*ocks, ofgj'aimlar jnicrocline mosaics included in the quartz.

110

Hex T. Piuder.

Plagioclase (10 to 15 per cent) is gonorally a slightly turbid and
poorly twinnod oligo-albite (AboAn^) which is partly replaced
by myrmekite when in contact witli microcline — it is never zoned,
and is invariably replacc'd in part by tine fibrous soricit(‘.

Biotite which, on the averages, forms 10 per cent of tliese rocks is
of two types — (1) A strongly pleochroic greenisli brown variety
with minute sagenitic rutile inclusions. The flakes are well
formed parallel to 001, but have ragged terminations. It is
partially replaced by green chlorite, especially along the cleav'age,
leading to an intergrowth of these two minerals. (2) A bright
greenisli completely chloritised biotite, usually developed in
(‘ontact with microcline and intergrown with epidote. The biotites.
often occur in small clustered aggregates- An the more crusht'd
varieties they occur in well defined bands, curving around the
lenticular granulated microcline aggregates.

Apatite is the most abundant accessory in stout idiomorpliic prisms.
Minor accessories are small zircons (with weakly pleochroic haloes)
included in biotite, scattered grains of magnetite and epidote.

Specimen No, 1213, an even-grained type, has been analysed as
typical of these granite gneisses, and the analysis appears in
Table 2, Column 1. It is a typical granite, and the appearance
of a little corundum in the norm is due to tlie slight secondary
alteration of the felspars. TJiere is insufficient excess alumina
to suggest the possibility of a sedimentary origin for tliese gneisses,
(Bastin, 1909, p. 401.)

TABLE 2.

Granite

Gneisses from the Toodyay Area.

1.

2.

SiOo

• . • • . .

71-85

63-28

TiO,

0-25

0-83

AhO,

15 • 00

14-56

0-55

3-13

FeO

... ...

1-20

5-10

MnO

Tr.

0-09

MgO

...

0-42

I -70

CaO

1-52

3-51

NaoO

3-67

3-66

K^O

4-45

2-58

HA)+

0-54

0-52

H,0-

0-02

0-15

0-11

0-45

99-58

99-56

Norms.

Q

29-52

20-40

Or

. • « » • •

26-13

15-57

Ab

• • • . ♦ .

30-92

30-92

An

... ...

6-67

15-01

C

1 -73

0-20

hy

2-58

9-74

mg

0-70

4-41

il

... ...

0-30

1-52

ap

...

0-34

1-01

1. Biotite-inicrocliiie granite gneiss (Spec. 1213), Toodyay, W.A. Analyst, K. T. Pridcr.

2. Biotite-oligoclase granite sneiss (Spec. 15389), Toodyay, W.A. Analyst. R. T. Prider.

Geology and Petrology of Part of Toodyay District, W.A. Ill

(ii) Microcline-granite gneiss.— -The mica is much less abundant than

in type (i). These rocks are rare and occur in narrow seams.
They consist essentially of microcline, quartz, plagioclase, and
accessory biotite, and, therefore, probably represent an originally
aplitic phase of the granite. The microcline is slightly micro-
pc^rthitic, and the less abundant plagioclase is a clear, finely-
twinned oligoclase (AbggAn^g). Such mica as is present is the
same greenish-brown partially chloritised variety as described
above.

(iii) Biotite-oligoclase-granite gneAss . — In this type, which is of rare oc-

currence, there is a much greater development of brownish biotite
than in (i) above. The main felspar is oligoclase, and microcline
is present to the extent of less than 10 per cent. The approxi-
mate mineralogical composition of a typical specimen (9577) is —

Quartz

20

volume

Oligoclase (Ab 4 An|)

65

%

Microcline ....

5

o/

0

Biotite

10

0/

0

Accessory rutile, zircon, and apatite.

The oligoclase is in subhedral crystals averaging 2 mm. diameter, with
oscillatory normal zoning and average composition Ab 4 Ani. Occasional
plates of microcline are present and, when in contact with oligoclase, myrme-
kite is developed.

The biotite is a brownish, non-chloritised variety, but still carries the
characteristic sagenitic rutile inclusions.

(6) XenoUths in the Uppe?‘ Granitic Gneiss.

(i) Schistose plagioclase amphibolites .- — These are found in narrow irregular
bands, varying from small stringers, several inches wide, which taper out
quickly, to well-defined bands up to one chain wide which usually run parallel
to the foliation of the gneiss. The occurrence in them of acid veinlets suggests
that they are inclusions, rather than sheet-like intrusions, in the gneiss, and
they often show a minute crumpling which accords with the view that they
are xenolithic, rather than post gneiss intrusions, as there have been no major
tectonic movements since the final consolidation of the granite gneiss.

There is little variation in these rocks and they are almost identical, both
mineralogically and chemically, with the plagioclase amphibolites, already
described, which occur interleaved with the meta-sediments. The main
points of difference may be summarised : —

(1) Xenoliths in the gneiss have hornblende with a much stronger

pleochroism : — X yellow-green ; Y deep olive green ; Z deep

bluish-green ; and absorption Y Z X.

(2) The plagioclase is more saussuritised and less abundant in the

xenoliths.

(3) Microline is 'absent from the xenoliths and quartz more abundant.

(4) Magnetite, rimmed with sphene, spread out into trains parallel to

the schistosity, is more abundant in the xenoliths.

A specimen of contorted hornblende schist (No. 1241) from a xenolith in
the granite gneiss was analysed and the result is seen in Table 1, Column 2.
The main points of difference from the previously described hornblende schists
lie in the lower Si02 and corresponding higher alumina in the xenolith. The

1]2

Rex T. Prider.

alkalies, especially potash, are also slighter lower. These features indicate
the close resemblance of the two rocks and also indicate that there has been
no addition of material to the xenoliths from the granite gneiss.

(ii) Hornhlendic schlieren . — These small lenses in the upper gneiss were
■only noted in one place (40 chains N., 527 chains W. from datum). They are
irregular shaped len tides up to 12 inches wide, which are elongated parallel
to the foliation of the enclosing gneiss.

That these schlieren are older than the intrusive granite gneiss is shown
by the narrow granitic veinlets penetrating them, and also by the fact that
small fragments of the xenolithic material are found in the enclosing gneiss.

The central }3ortion of these inclusions is made up largely of a dark green,
bladed hornblende, in crystals up to 0-5 cm. long, with occasional larger
plates of a pale greenish pyroxene. Small white angular patches and veinlets
of felsjDathic material are present. Near the edge of the xenoliths the rock
becomes lighter in colour, hornblende becomes rarer, a pale greenish pyroxene
taking its ]>lace. The enclosing gneiss is a fine granulitic microcline granite
gneiss.

Under the microscope the fine microcline gneiss is seen to have pale
gi*eenish liornblende derived from the hornblendic xenolith, as its ferromag-
nesian. Approaching the contact the most noticeable features are (1) the
increase in the hornblende content, (2) entrance of idioblastic sphene and
colourless pyroxene, (3) decrease of microcline and corresponding increase
of oligoclase. The felspars at the actual contact are considerably epidotised.

The xenolith itself is made up mainly of pyroxene, hornblende, and
introdiiced quartz-oligoclase veinlets. The outer zone of the xenolith is
rich in pyroxene (in large plates up to 4 or 5 mm. diameter), which under the
microscope are seen to be fringed with pale greenish hornblende. The pyroxene
in thin section is colourless, with 110 cleavages well developed and a marked
parting ]>arallel to 100 and less j^erfect parting on 010.

The properties Z a c = 42° ; y = 1-705 ; « = 1*676 ; y - a =
•029 ; biaxial (+), and very weak dispersion, point to a diopsidic pyroxene
with 20 per cent, of the hedenbergite molecule.

The amphibole which is clearly developing from the ])yroxene is a pale
greenish, somewhat fibrous va^-iety, with an irregular coloration in the same
crystal. Pleochroism is X yellow-green ; Y olive green ; Z bluish-green ;
and absorption X<Z<Y. Z A c ^ 23°. This is a common blue-green horn-
blende. It is usually developed where the pyroxene is in contact with oligo-
clase and rarely where in contact with quartz, the hornblende having apjmrently
obtained its alumina from the felspar.

In the centre of the xenolith, although most of the diopside has been
replaced by amphibole, cores of diopside in this mineral [)oint clearly to its
origin.

These xenoliths have been originally dio]}sidic rocks, now converted into
rocks composed ]3redominantly of hornblende. An anomalous feature is that
the cores show considerably more alteration than the periphery. There has
been a slight hybridisation of the adjoining granite (gneiss) giving rise to a
fine-grained hornblende granite gneiss. This hybridisation of the granite is
developed on a larger scale in the Lower Gneiss and this ciuestion will I)e
considered more fully at a later stage.

(iii) Biotite granidUe xenoliths . — These are much finer-grained and carry
more ferromagnesians than the enclosing gneiss, in which they occur as
irregular shaped elongated darker-coloured patches, in which the foliation
approximates to that in the surrounding gneiss. All these rocks are alike

Geology and Petrology of Part of Toodyay District, W.A. 113

petrologically, being composed of partly chloritised biotite, quartz, and oligo-
clase with accessory magnetite, apatite, and sphene. One specimen, however,
shows a development of a deeply-coloured hornblende.

The boundaries of the xenoliths are sharp and there is no transitional
hybrid zone. The structure is fine, even granular, schistose (figure 12). The
plagioclase carries minute rodlike inclusions of recrystallised muscovite. It
is rarely twinned but its refractive index, approximately equal to that of
Canada balsam, indicates that it is an oligoclase.

Biotite-quartz-plagioclase graiiulite xenoliths in granite gneiss.

A. No. 1220. Edge of xenolitli. The left hand edge of the
field is occupied by the coarse granite gneiss, the right hand por-
tion by the biotite-quartz-]3lagioclase granulite.

B. No. 1221. Variety with deep greenish hornblende.

The biotite is a greenish-brown variety, altered along cleavages to chlorite.

The deep green hornblende present in one of these rocks tends to occur in
small clots, often enclosing magnetite.

Accessories are apatite, magnetite, epidote, and sphene.

These rocks represent psammitic types of sediments, which with the
admixture of a little calcareous material have at times developed hornblende
and epidote ; they cannot be matched with any of the metasedimentary
rocks in the Jimperding Series.

(iv) Eulysitic rocks (meta-banded ironstones). — Banded quartz-magnetite-
grimerite rocks with affinities to the eulysites have been noted vas enclosures
in the Upper Gneiss in t-wo places : (a) At 26 chains E., 40 chains N. of datum.
This occurrence is in the form of a band, several chains wide, which can be
traced for about 30 chains in a south-easterly direction. (6) At 92 chains W.,
140 chains N. of datum. This occurrence is also a band in the gneiss, about
1 chain wide and can be traced for about 30 chains in a north-easterly direction.
Both of these bands lie close to the bottom of the Upper Gneiss and are
probably both part of the same horizon. The rocks from both occurrences
are essentially the same, being magnetic banded rocks composed of varying

114

Rex T. Prider.

proportions of (juartz, magnetite, and grunerite. These rocks are similar
to some occurring as xenoliths in the Lower Gneiss, which are described in a
later section ((5) (e) (i)) of this paper.

(c) Sunwiari/ of Cone/ unions regarding the Origin of the Upper Gneiss.

The main mass is constant throughout and is a normal hiotite-microcline
augen gneiss, derived from a ])orphyritic microcline granite. The presence
of both st'dirnentary and igneous xenoliths and the occurrence of small trans-
gressive apo])hyses in the underlying ([uartzite, ])oint clearly to its intrusive
<'haracter.

Tlie gneissic banding was developed when the rocks had partially crystal-
lised (/.e., it is a fluxional foliation due to the alignment of the microcline),
during which time {jrotoelastic structures were developed. The final crystal-
lisation of ([iiartz took place after the tectonic movements had ceased.

The microcline granite was intruded as a thick sill which has produced no
a|)])arent thermal effects in tlie associated sediments which are in every instance
(juartzites. Such rocks, however, would not be expected to yield any in-
formation regarding the degree of thermal metamorphism induced by the
granite. Felspathisation of quartzose rocks, about which a good deal has
been written, is not developed in the Toodyay Area. It is not clear why the
<juartzites al)ove this gneiss have different microstructures from those below
the gneiss. If the granite we're a sill we should expect them to be identical.

[d] The Lower Granite Gneisses.

These rocks, outcropping in the north and north-east parts of the aiva,
^'arv from tine, even-grained, well-banded tyjDes, to coarse porphyritic granites
in which the banding is visible only in a flow oritmtation of the microcline
phenocrysts.

The well-baiuU'd varieties all have granoblastic structure with but little
sign of granulation. A gneissic structure is evident in the sub-])arallel orienta-
tion of the biotite and in the elongation of the felspar and quartz allotrio-
mor])hs in a common direction. In these finer-grained gneisses the (piartz
grains have an average index of elongation of about 3:1.

The mineralogical composition of the Lower Gneisses is fairly constant,
irrespective of whether the rock is fine-grained or coarse por[)hyritic. Tlie
av('rage composition by volume is : - -

Oligoclase (Ab^Aiii) ....

30°o

Microcline (sliglitly micropcrthitic) ....

30%

Quartz ....

30“„

Biotite ....

I0°o

with accessory apatite, magnetite, and more rarely sphene and e[adote.

Minor variations due to a greater abundance of biotite together with an
increase of the pro],)ortion of oligoclase to microcline (as in the Upper (ineiss)
have been noted. Also a hornblende bearing granite gneiss is develoj)ed by
liybridisation of the granite* by the hornblendic xenoliths which are present in
some abundance.

The main point of difference from the Upper Gneiss lies in the general
absence of eataclastic (or protoclastic) structures.

The microcline ])henocrysts which in the Upper Gneiss are represented
by lenticular "eyes” are he*re seen to be well-shaped, uncrushed crystals (up
to I inch in length).

Gp^ology and Petrology oi’ Part of Toodyay District, W.A. 115

Biotite-oligoclase gneisses similar to those in the Upper Gneiss have
been noted in narrow bands. Spec. No. 15389 is a type containing approxi-
mately 25 per cent, of a brownish biotite. It has been analysed as typi(^al
of these rocks (Table 2, No. 2). It differs from the microcline bearing gneiss
in its higher iron and lower silica and ])otash (content.

Under the microscope, oligoclase in subhedral grains up to 4 or 5 mm.
<liameter is the dominant constituent. It shows line albite twhming, the
lamellae often being slightly ciirved due to strain. A noticeable feature is that
the (piartz moulded on such a strained crystal shows no undulose extinction.
The brovv-nish biotite is in flaky aggregates winding around the Iarge?r oligo-
clases ; they feather out against one another and have V>een squeezcnl together
by movements in the semi-cTystallised magma. The microstructurcs recall
in many ways the round grained ” gneisses of Glen Doll, Forfarshire (Ilarktir,
1932, p. 298).

Aj>atite is a very abundant ac.cessory, in small stout [jrisms, closely
associated with the biotite ricli bands.

These biotite-oligoclase granite gneisses a])])ear to be tlie result of the
crystallisation of the residuum squeezed off! from the (uxrlier formed microcline,
tlms containing a (H^ncentrate of biotite, oligoclase, and apatite.

(e) XenoUlhs in the Lower Granite Gneiss.

Numerous oval areas of foreign enclosures varying from several yards
in diameter, to elongated masses up to 10 (diains in kmgth have been noted
in the i^owor Gneiss. They are most abundant in the eastern part of the
area (Plate 1). Most are poorly exposed and, witli several excej)tions, the
contacts with the enclosing gneiss are obscured by soil. The mai>j)ing, hoW“
ever, indicates that they are lenticular bodies completely surrounded by
granite gneiss. Tlie amphibolite enclosures are frequently traversed by
quartz-felspar veins coming from the granite gneiss, thus establishing their
pre-gneiss age. The longer axes and foliation of the entdosures strike jjaralle)
to the banding in the surrounding gneiss. There is considerable variety
amongst these xenoliths and the main ty[)es will be considered separately
as follows : —

(i) Kulysitic rocks.- Thono rocks are scarce in this area, but their occur-
rciico is of interest since they boar similarities to rocks occurring as bands
in the Bolgart greenstones to the n<jrth of Toodyay, and sinc^e they are rep-
res(uitatives of the somewhat rare eulysites.

They are found in a small xenolith 10 chains 8., 223 chains F. of datum
(Plate I), entirely surrounded by gneiss. Near}>y xcmoliths are greenisli
“ hornblenditos.”

'^wo rocks were collect(;d from this locality and in view of their somewhat
dif£(H’cnt cliara(T(T they will be described so])arately.

Jianded (luartz- magnetite- hyper sthene rock (8'pec. No. 1 5451 ).

This is a heavy, (^oarsedy- banded typ(j made u[j of bands (3 mm. wide) of
silky lustr(‘d lamellar hyi)ersthene, alternating with darker magnetite bands
fi'om 1 to 2 mm. wide. The rock is (^onsi(lerably weatliertid and iron stained,
and its feeble magnetic (diaracter indicatt^s that th(i magnetite lias gone largely
to fine granular martite.

Th(j hypiTstliene is usually found with its longitudinal (d(;avago at about
45° to the banding. It has a poikiloblastic structure, enclosing botli (piartz
and magnetite, and the original bedding has been preserved in the wide bands
of magnetite and the thin |)arallel bands of fine granular ore which traverse

116

Rex T. Prider.

the hypersthene (figure 13B). The hyperstheiie show>s a fine lamellar structure,
(’leavage is poorly developed, but there is a good j^arting on 010. Although
l>rownish (due to slight weathering), the weak j)leochroism is still visible.
Its characters— X a ; Y = b ; Z = c ; (-)2V 83° ; y 1 -765 ; (*

•--- 1 -74-5 — indicate an iron rich hypersthene with apjjroximately 85 |.er cent
of the orthoferrosilite molecule (Henry, 1935, p. 223). (‘omparing the above
data with those given by Henry (p. 223) for tlie iron rich hypersthenes, it
will be seen that this hypersthene agrees almost exactly with the data given
for the hypersthene in the eulysite from Mansjo, Sweden, which lias (-) 2\’ =-
83° ; y == 1-769 ; a 1-751. This is the most iron rich tyjie described
])y Henry (containing 44-93 per cent FeO).

Pig. 13.

Banded eulysitie rocks.

A. (^uart/v-lienuitite-grunorite rock (No. 17452).

B. (,)uart/i iimguetlte-liypersthene rock, showing original bed-
ding preserved in the parallel trains of small magnetite grains
included in hypersthene. The hy])ersthene is altering along its
edges and along irregular cracks to a fine fibrous ferro-nntho-
])hyllite.

The hypersthene is altering to a more fibrous, highly birefringent am] hi-
liole, both around its edges and along irregular clacks. Some of the hyper-
sthenes are eompletely replac'ed by tliis material. The optics of this ami'hibole
are Z ^ c ; (-) 2V large ; y 1 -687, and it is therefore a ferro-anthojibyllite
with apjiroxirnately 60 per cent FeSi 03 .

Quartz and magnetite are the only other constituents with the exception
of rare apatite. Both are closely associated and there is no sign of reaction
betw'een them. The magnetite is in liands up to 2 mm. wide and in trains
of small grains (parallel to the bedding) included in hypersthene. In ]daces.
tongues of magnetite connect successive bands ol iron ore. The almost
non-magnetic character of the rock indicates that the magnetite lurs gone
to fine granular martite, and this is confirmed by the cherry red streak of
tlie dark bands. On the outer portion of tlie rock the reddisli lu'matite can
be seen replacing the magnetite, and the change throughout the rock is
ascribed to oxidation due to weathering. A very different occurreuee of
hematite is seen in the other specimen from this xenolith which is doserihod
bc'low.

Geology and Petrology of Part of Toodyay District, W.A. 117

The mineralogical composition (by volume) of the hyperstheiie rock
is approximately —

Hypersthene .... .... .... 60 %

Ferro-anthophyllite
Martite (after magnetite)

Quartz
Apatite

15 %

15 -

-20

5 %

Accessory.

Banded quaTtz-hematite-grunerite rock (15452).

This specimen is from the same locality as (a) above — the field relations
lietween the two types are obscure.

It is a finely banded, even-granular rock, and consists of alternating
dark coloured bands of recrystallised hematite and pale greenish ([uartz-
amphibole bands varying in thickness from 0-5 to 1-0 mm. The sj^ecimen
is non-magnetic — this, together with the deep red streak confirms tlie de-
termination of the silvery grey, metallic mineral as hematite.

Under the miscroscope the structure is even-grained granoblastic gneissic,
the hematite 0''*f‘urring in recrystallised grains and aggregates showing a
marked segregation into bands (figure 13A) — quartz is often included in
these aggregates and shows no signs of reaction- with the liematite.

The amphibole and (piartz form a granoblastic aggregate, the latter
often occurring as poikiloblastic inclusions in the former. The amphiboles
have a random orientation in the rock, although in ])articular bands they
tend to show the same orientation througliout.

This a-mpliibole is a })ale greenisli weakly pleochroic variety- in irregular
shaped prisms with lamellar structure. Tlie ]:>rism faces are fairly well
developed, but the terminations, whether in contact with (piartz or iron ore,
are irregular. It has good 1 10 cleavage and commonly shows multiple
twinning on 100. The oj^tics are a -= 1 -635 ; y ^ 1 -660 ; y - a ^ *025 ;
Z Ac — 15° ; (-)2V near 90. Opt. ax. pi. || 010.

The refractive indices indicate a cummingtonite with 42 per cent
FeSi03, but the negative optical cliaracter suggests admixture of the actino-
lite molecule. Richarz (1927, p. 700) has described a similar amphibole

from the Lake Superior District. It has Zac = 15° ; y = 1-680 •

/3 = 1 • 668 ;

a = 1 • 665 ;

y - a *025 ; (-)2V near 90° and com-

position : —

AI2O3

5-3%

Fe^Oa

00

0

0

FeO

22-4%

CaO

•> ■ gC/

- ^ 0

MgO

a • 4 c/

thus differing

from grunerite

in its high FegO^ and Al^Og (nntent. In its

optics it closely resembles the Toodyay mineral whicli is probably an alum-
inous type of grunerite with admixture of actinolite molecules.

The origin of the banded- eulysitic types.

The [ireservation of well marked banding in these rocks is indicative
of their origin from sedimentary, bedded iron ores.

In their appreciable MgO content, and, in the case of type (b), the
possible presence of CaO, the original rocks would appear to liave been impure
banded siliceous ankerite rocks rather than greenalites. In the hypersthene-

118

Rex T. Prider.

quartz-magnetite rocks, the original carbonate must have been largely siderite
with but small admixture of magnesite. The metamorphism has been
effected under thermal, rather than regional conditions, as evidenced by
the complete lack of orientation in the amphiboles and orthopyroxenes formed
dining this period.

In the case of the quartz-hematite-grunerite rock {&), it would appear
that limonite was a constituent of the original quartz-carbonate sediment,
and has, by re-crystallisation given rise to well crystallised hematite without
any reaction with the quartz. Si02 has reacted only with the (Fe, Mg)
carbonates to give rise to the grunerite. Such a quartz-limonite-iron car-
bonate rock is known amongst modern bog iron deposits (Van Bemmelen,
1900, p. 319).

The presence of considerable MgO and possibly CaO, and the variable
proportions of quartz, iron ore, and amphibole (or hypersthene) throughout
the rock seems to indicate that greenalite cherts were not the sediments from
wliich the eulysitic rocks were derived, and while the sedimentary origin
is clear, there is some doubt regarding the original character of the sediment,
but it appears most probably to have been a banded limonite-iron carbonate
rock.

(ii) Calc-silicate rocks . — Xenoliths of this type have only been noted
in one place (114 chains X., 113 chains E. from datum). The rocks form
a well defined band in the coarse porphyritic gneiss, rimning parallel to the
strike of the enclosing rocks. The lime silicate rocks occur in bands and
lenticles running through a band of white vein-like quartz.

The rock is similar to specimen 1249 (described earlier), differing only
in the relative proportions of diopside and grossularite, and in being more
siliceous. It has the following approximate mineralogical composition
(Vol. %)

Quartz 50 per cent, diopside 18 per cent, epidote 5 per cent, grossu-
larite 25 per cent, amphibole and sjihene 2 per cent.

Isotropic grossularite is the only idioblastic mineral, and it encloses
diopside and occasionally quartz.

Diopside, which is not enclosed by grossularite, is altering to a fine
fibrous colourless amphibole.

These rocks undoubtedly have the same origin as the lenticles of lime
silicate rocks in the lower quartzites. They represent xenoliths of the
Jimperding metasediments which have been caught up in the intrusive
porphyritic microcline granite.

(iii) Co7'dierite-anthophyllite rocks and related types , — -These rocks are
found in a large xenolith in the granite gneiss at a position 236 chains E.,
177 chains S. from datum. The occurrence is more or less circular in shape
and about 5 chains in diameter. It consists largely of anthophyllite-hyper-
sthene-pleonaste rocks, but several other types, viz., cordierite-anthophyllite
and biotite-clinochlore-anthophyllite rocks are found here.

These rocks have been fully described elsewhere (Prider, 1940), but
the cliemical analyses are repeated here in Table 3. The conclusion re-
garding the origin of these anthophyllite-hypersthene-spinel rocks is that
they \\'erc‘ derived from a hypersthenite magma contaminated by aluminous
material. The cordierite-anthophyllite assemblages were developed from
the spinel hypersthenites during the period of intrusion of the granite gneiss,
by the addition of silica from the granite magma.

Geology and Petrology of Part of Toodyay District, W.A. 119

TABLE 3.

Analyses of cordierite-anthophylUte rocks and related types from Toodyay, W.A. {quoted

from Prider, 1940,

p, 377).

1 .

2 .

3.

4.

SiO.,

30-91

30-83

33-20

49-73

21-36

20-47

19-75

12-70

FegOg

/19-971

9-23

3-36

4-56

FeO

\ /

11-28

13-23

12-27

MgO

23-57

16-10

21-57

16 -.59

CaO

Tr.

Tr.

Nil

Tr.

NagO

Nil

0-36

0-24

0-40

K^O

Nil

5-18

0-22

0-54

HgO+ ...

2-58

4-87

8-43

2-77

H^O- ...

0-90

0-42

0-10

0-12

TiOa

1-66

0-70

0-06

0-21

P 2 O 5

Tr.

nd.

nd.

Tr.

MnO

0-16

0-07

0-14

Nil

Cr^Oa ...

0-17

Nil

Nil

Nil

101-28

99-51

100-30

99-89

1. Olivine-spinel-anthophyllite-hypersthene rock, Toodyay, W.A.

2. Biotite-clinochlore-magnetite-corundum rock, Toodyay, W.A.

3. Corundum-spinel-anthophyllite-cordierite-clinochlore rock, Toodyay, W.A.

4. Biotite-cordierite-anthophyllite rock, Toodyay, W.A.

(iv) The Amphibolite Xenoliths. — Amphibolites, varying from almost
pure hornblende rocks to quartz-plagioclase amphibolites in which the
felsic minerals are in excess of hornblende, are the most abundant xenolithic
types in the lower gneisses. They are found in lenticular masses, measuring
up to 10 chains long x 3 chains wide, and in thinner bands which may be
traced for greater distances. The elongation and lamination in the amphibo-
lites are parallel to the strike of the enclosing gneiss. The occasional pre-
sence of narrow sill-like veinlets from the granite gneiss at the edges of the
amphibolite masses indicates the intrusive character of the gneiss. There
has, in some instances, been a considerable hybridisation of the granite
gneiss, which has been changed from its normal character into a more basic
hornblende granite gneiss.

These amphibolites comprise a number of types which are illustrated
by the following description of typical members : —

Quartz-plagioclase amphibolites.

These vary from medium-grained dark coloured rocks with a visible
lamination (15438) . to types with a more granular structure in which no
lamination is visible (e.g., 15444).

The former are darker and more granular than the hornblende schists
described earlier in this paper. The non-foliated granular type of amphibolite
is the most common, and No. 15444 will be described as typical of this group.
In hand-specimen it is a dark coloured granular rock with only a slight trace
of parallelism of the hornblende which is the dominant constituent. It is
a dark green variety in well shaped prisms 1 mm. in length, the interspaces
between these prisms being occupied by light coloured fine granular felsic
material.

Under the microscope the structure is granoblastic, and the rock is made
up of brown-green hornblende (65 per cent), saussuritised plagioclase (15
per cent), clear oligoclase-andesine (10 per cent), cpiartz (10 per cent), with

120

Rex T. Prider.

accessory epidote, magnetite, and sphene. The amphibole is idioblastic
(figure 14A), in prismatic forms up to 2 mm. in length, usually clustered
together, these groups being separated by finer granoblastic tpiartz-plagioclase
aggregates.

Some of the hornblende has a poikiloblastic structure carrying rounded
inclusions of cpiartz, magnetite, and, more rarely, plagioclase. It is a much
deeper coloured variety than that developed in the hornblende schists inter-
bedded with the quartzites, and has pleochroism : — X yellow-green ; Y dark
olive gret‘ii ; Z deeji bluish green ; and absorption X < Y Z, ^ = 1-675.
Roth (piartz and plagioclase are xenoblastic. Two varieties of the latter
are present in ajqH’oximately ecpial amoLint (1) a completely saussuritised
variety, and (2) a clear rarely twinned variety often showing slight zoning.
It is an oligoclase-andesine near Ab 7 Au 3 .

Fig. U.

Amphibolite xenoliths in the Lower Granite Gneiss.

A. Medium grained amphibolite (15444). Con-

stituents are: hornblende, turbid plagioclase, quartz, magnetite
with rims of sphene.

B. Coarser amphibolite (15441), showing poikiloblastic in-
clusions of quartz and magnetite in hornblende.

C'. ‘ ‘ lIornl)lendite ’ ' — Hare jioikiloblastic inclusions of quartz

in the hornblende.

Magnetite is the most abundant accessory and is invariably rimmed
with splume indicating its origin from ilmenite. Epidote and apatite are
rarer af*cessories.

In coarser grained varieties (e.g., 15441), the felspar is completely re-
placed by a fine granular sericite-opidote aggregate, and the larger, more
abundant hornblende plates show' a well developed poikiloblastic structure
(figure 14R) with inclusions of quartz and magnetite, the latter often with
a narrow rim of yellow’ highly birefringent epidote.

Hornblendites.''

With a decrcas(‘ of the quartz-felspar content the C|uartz-plagioclase
amphibolites pass into almost pure hornblende rocks in wliich the only other
constituent is an occasional small grain of quartz. 15436 is an example —
ill hand speudmen it is made up of a granular aggregate of hornblende (figure
14C), noticeably lighter in colour than in the above types. Under the

Geology and Petrology of Part of Toodyay District, W.A. 121

microscope many of the hornblende prisms carry small rounded poikiloblastic
inclusions of clear quartz. Rare felspar is evidenced by the occurrence of
small patches of fine granular epidote.

Pyroxene-plagioclase amphibolites.

Tliese rocks are similar to the (piartz-plagioclase amphibolites with the
addition of a pale greenish diopside. They are like many of the hornblendic
rocks of the Lewisian of Scotland.

No. 15445 is typical (figure lOA). Tt is coarse, even-grained, granoblastic,
Avith no tendency to gneissic structure. The constituents are hornblende
{(iO j)er cent), diopside (25 per cent) and oligoclase (15 per cent), with
a'-C(‘SSory tpiartz and apatite.

The hornblende, in plates to 3 mm. diameter, sometimes moulded arormd
iiiopside, is a. brownish-green variety similar to that in the quartz-plagioclase
amphibolite. Tt is noticeably paler in colour at its junction with the diopside.
Poikiloblastic quartz inclusions are common.

The pyroxene is a pale greenish diopside with Zac = 38°. It has a
curious mottled appearance due to small platy inclusions of pale green horn-
l)lende, which appear to be developing at the expense of the diopside.

The fels])ar is an oligoclase (Ab^An^) with fine albite twinning, occurring
in subhedral to anhedral grains which are, in plac'es, associated with horn-
blende in a sub-ophitic fashion.

Apatite, in euhedra averaging 0*2 mm, is the most abundant accessory.
Quartz is very rare.

Chemical Composition of the Amphibolites.

ypecimen 15441 (described above) was analysed as representative of
the most widespread type. The analysis appears in Table 1, column 3.
The similarity to the schistose plagioclase amphibolites 1 and 2 is at once
evident. The igneous character of these rocks has been noted on an earlier
l^age and need not be considered further here. Except for its slightly higher
alumina and low lime and magnesia, this amphibolite analysis agrees verv
<'losely with that of the later dolerites (Table 1, col. 4). from this area.

The am])hibolites, then, appear to be derived from basic igneous rocks
a])proximating to dolerites or quartz dolerites in composition. Tilley (1921,
pp. 98-116), Avho has described a number of amphibolite enclosures in the
granite gneisses of the Southern Eyi-e Peninsula, South Australia, which are,
in many ways, similar to those found at Toodyay, has outlined (pp. 108-109)
the chemistry of the conversion of pyroxene to amphibole and little would
be gaiiK'd by repeating this here.

Wliere the hornblende becomes more abundant in these xenoliths its
crj'stals beconn^ larger and apparently more aluminous at the expense of the
I>lagioclase. Quartz persists, as a by-product of the amphibolitisation, even
in the almost pure hornblendic types.

The pyroxene-plagioclase amphibolite xenoliths appear to repres(mt
completely recrystallised older basic rocks, such as occur in sill-like bodies
of pre-tectonic age, interbedded with the Jimperding metasediments. The
grain of these rocks has become coarser as a result of the recrystallisation of
tlie amjdiibole. Basic xenoliths are often of fine grain (Joplin, 1935, p. 233)
ascribed in part to the disruption of highly poikilitic crystals of hornblende.
In the Toodyay gneisses the amphibolites have not suffered any such breaking
up of the hornblende.

122

Rex T. Prider.

(/) Hybridisation of the granite g^ieiss.

There is a considerable development of quartz-oligoclase-hornblende
gneisses in the Lower Gneiss as a result of hybridisation of the granite by
basic inclusions. These types are well developed 80 chains N., 300 chains
of datum, where the lower gneiss is seen to transgress the bedding of the
hornblende schists and metasediments. The gneisses are of variable com-
position averaging : —

Hornblende

Oligoclase

Quartz

Chloritised biotite

30 to 35 per cent (by volume).
45 to 50 per cent.

15 per cent.

5 to 10 per cent.

Microcline is rare and the accessories are : magnetite, sphene, epidote, and
^patite. The plagioclase is usually much saussuritised but when determinable
s an oligoclase (Ab^An^). The hornblende is a slightly deeper coloured variety
han in the associated schists, and the biotite is the characteristic greenish

Pig. 15.

Granite gneiss — amphibolite hybrids.

A. 15640. Showing clotted aggregates of hornblende (often
with poikiloblastie structure). The other constituents are turbid
plagioclase and clear quartz.

B. 15397. Hornblende granite gneiss. Constituents are
deeply coloured hornblende, somewhat fibrous pale greenish
ainphibole, oligoclase, quartz, idioblastic sphene (with central ore
inclusions) and apatite.

chloritised variety, with sagenitic rutile inclusions. The hornblendes are
often clustered together (fig. 15A) and may have a poikiloblastie structure.
In specimen 10420, lenticular remnants of schistose plagioclase amphibolite
are present in a hornblende-granite gneiss.

A hornblende-granite gneiss (15397) from the vicinity of a coarse amphi-
bolite enclosure has been analysed (Table 4, No. 2) and will be described in some
detail. It is an even granular rock -with a gneissic structure and it contains
several sill-like quartz-felspar layers.

Geology and Petrology or Part or Toodyay District, W.A. 123

Under the microscope the constituents observed were : blue-green horn-
blende (20 per cent), oligoclase (55 per cent), quartz (20 per cent) with
diopside, sphene, magnetite, apatite, and microcline as accessories. The struc-
ture is granoblastic with no tendency to parallel alignment of the constituents.

Hornblende, usually idioblastic, occurring in small lenticular clots is, in
places, altered to a paler greenish fibrous amphibole. Xenoblastic oligoclase
(AbgyAuig), in slightly turbid grains averaging 1 mm. diameter, is the only
felspar — it occurs in granoblastic aggregates with quartz and is frequently
elongated slightly, parallel to the banding. The absence of microcline is un-
usual, but is explained by the fact that the microcline in the granite gneiss
has crystallised early, forming large phenocrysts, and that specimen 15397
is the result of the reaction between the residuum and the basic xenoliths.
Apatite, magnetite, and sphene aj-e all abundant accessories ; the sphene is
usually idioblastic and carries central inclusions of magnetite (fig. 15B).

The analysis of this rock, together with that of the microcline granite gneiss
and coarse plagioclase amphibolite appears in Table 4. It is clearly inter-
mediate between the other two t3q>es. The main discrepancy is in the alkalis,
but this, as noted above, is due to the removal of the microcline phenocrysts
from the magma prior to its reaction with the amphibolite.

TABLE 4.

Aymh/sis of Hornblende Granite Gneiss {Col. 2) compared with Coarse Plagioclase
Amphibolite (1) and Microcline Granite Gneiss (2).

SiOo

TiO;

AI2O3

Fe,03

FeO

MgO

CaO

NagO

K2O

H3O+

H^O-

MnO

P2O5

FeSa

1.

2.

3.

49-05

64-16

71-85

1-07

0-74

0-25

15-03

15-12

15-00

3-16

1-95

0-55

9-08

2-90

1-20

6-96

2-13

0-42

10-47

6-63

1-52

1-70

4-48

3-67

0-95

0-20

4-45

1-43

0-67

0-54

0-25

0-04

0-02

0-09

0-18

trace

0-08

0-32

0-11

0-13

0-15

nd.

Total

99-45 99-67 99-58

1. Coarse amphibolite xenolith (refer Table 1, No. 3), Toodyay.

2. Hornblende granite gneiss (No. 15397), Toodyay. Analyst, R. T. Prider.

3. Biotite-microcline granite gneiss (Table 2, No. 1).

Much more work, both field and laboratory, is required before any definite
statement can be made regarding the origin of these acid hornblende bearing
gneisses. So far as examined, however, they appear to represent a hybridisa-
tion of the intrusive granite gneiss.

(g) Summary of the conclusions regarding the origin of the Lower Granite
Gneiss and the associated xenoliths.

The gneiss is essentially the same as the upper band but has not suffered
such granulation — it is essentially a porphyritic granite, with flow orientation
of the microcline phenocrysts. A less abundant biotite-oligoclase gneiss is con-
sidered to represent a more sodic phase of the main mass, which crystallised from
residuum squeezed out from the earlier phenocrystal microcline. Late stage
veins of garnet-aplite and pegmatite are developed from this magma.

124

Rkx T. Prider.

Xonoliths of various types are described : —

(1) Eulysitic types developed from banded iron ores, probably sideritic

and other carbonate types.

(2) Grossularite-diopside-qiiartz rocks derived from impure argillaceous

limestones.

(3) C’ordierite-anthophyllite rocks and relate*d types deri\'ed from

ultrabasic igneous rocks.

(4) Amphibolites, resulting froTU the reconstitution of basic igneous

1 ‘ocks.

\^.~ --The Younger Igneous Intruslves.

(1) The Granites.

The later non-foliated granites are confined to the south-west part of
the area when* tliey are intrusive into the series described above. The gran-
ites are cut by later pegmatites, aplites, and cpiartz veins, which often pene-
trate the nearby metasediments and which are considered to be the source
of the auriferous deposits found in the mica schists close to the granite.

The granites are always medium to coarse even-grained, remaining coarse-
grained right to their contact with the metamorj^hic rocks, indicating that
intrusion took place at some depth into already heated sediments. There
is no sign of the porphyritic and foliated structures seen in the older granites
occurring as bands in the Jimperding Series.

The later granitic intrusions are represented by thrf'e phases, which may
may be described briefly as follows : —

(u) The )iormal granites are coars(* textured rocks consisting of (piartz,
slightly perthitie microcline, sericitised oligoclase (Ab 4 Aii 2 , often
with zonal alteration), abundant myrmekite, irregular chloritised
biotite flakes with intergrowths of epidote, and accessory apatite
and sphene. In their mint'ralogical composition these rocks
are \'ery similar to the older granite gneisses described above.
(6) Garnet-nniscovite granites are even, fine to medium grained, allo-
triomorphic granular structured rocks made up of quartz, slightly
perthitie microcline, oligoclase and muscovite, with accessory
pink garnet {altered along irregular cracks to greenish biotite)
and a little brownish biotit(\ These represent a hypabyssal phase
of the granite and occur as dykes in the metamorphies close to
the main granite mass.

(c) Pegntatites, garnet aplites, and later quartz veins represent the final
phase of the granite intrusion. They are found in dykes and
\eins in botli the granite and adjacent metamorphies. The peg-
matites are coars(* grained microcline and muscovite bearing types,
in which the presence of molybdenite, columbite, and beryl have
been noted- — the two latter by Simpson (personal communication).

The garnet aplites are the tine grained equivalent of the garnet-
museovite granit(*s. The prevailing texture is fine equigranular
granitic, and the constituents are quartz, microcline, and oligo-
clase with accessory biotite (rare) and small pink gai'uets. The
aplites often form a part of the pegmatite veins.

WJiite quartz veins, representing the final ultra-acid ])hast* of the
granite, are fairly numerous in the metamorphies close to the
granite. A flat di]3ping f[uartz vein in a roof pendant of mica
schist in the granite has been proved to be auriferous, a test parcel
(50 tons) of this ore mined several years ago yielding 15dwt.
gold per ton.

Geology and Petrology of Part of Toodyay District, W.A. 125

{2) The Greenstones.

(a) Quarts, dolerites.- -The^se are the latest intrusives into the Jimperding
.Sei'ies. They occur as dykes up to five chains wide, most of which trend a
little \V. of N., although there are several large dykes with an E.-W. trend.
The geological map (Plate I.) shows the distribution of these dykes. They
all belong to the same period of intrusion and no examples of one dyke being
cut by another were found. This period of intrusion was much later than
the last orogenic movements in this region and the doleritic rocks show no
alteration other than those of deuteric character.

They are variable in grain from line basaltic to coarse gabbroidal, and
■r)phitic texture is characteristic except where obscured by extensive uralitisa-
lion of the pyroxene in the more acidic types.

Pig. 16 .

Quartz dolerifes

A. Quarts rare. Both pyroxene and plagiodase are un-
altered. The ophitic texture is well marked. The i)vroxene
cleavages are slightly curved and incipient fracturing is notice-
able. A little primary brown green hornblende is seen in upper
right in the vicinity of the end pbase fpiaitz. The iron ore is
ihnenite.

B. End phase of the (juartz <lolerile magma showing a con-
centration of quartz and apatite. Pibrous uralite replaces
pyroxene (ui)])er left) and the idiomor])hic plagiodase is highly
saussuritised. The ferromagnesian is mainly a brownish prim-
ary” hornblemle.

Th(‘ rocks vary from cjuartz-frc(' doh'ritcs made up of slightly browiiisli
])vroxeuc and plagiodase (Ab;|Anj) with apatite and ilmenite as accessories,
through (juartz dolerites which usually show considerable uralitisation of
the pyroxene, to more acidic ty])cs made u]) larg<dy of plagiodase, uralitised
])yroxf‘ne, ])rimary brown-gr(‘c‘n hornblende, and quartz, with leucoxenised
ilnnuiite and apatite as abundant accessories. The uralitisation of the pyrox-
eii(‘ is definitely a deuteric proce.ss in these rocks because there are no later in-
trusives or ('arth movemt'nts which could have effected thes(‘ changes, because
it is developed much niort* exttuisively in types in which the end stage (fuartz
1 as b('(‘n concentrated, and also bc'cause both unaltered and highly uralitised
rocks occur in the sarru' dyk('.

12(3

Rex T. Prider.

The microscopic characters of c(uartz dolerites have been so frecpiently
and fidly described that little space will be devoted to the description of the
Toodyay quartz dolerites which do not differ in any way from the normal
type of this rock. The only variations are in the proportion of micropeg-
ma.tite and the degree of deuteric alteration.

The minerals developed in these rocks are : — ■

Pyroxene, which is always a pale brownish, non-pleochroic, monoclinic
variety, of subhedral to euhedral development, usually penetrated opliitically
by plagioclase. There is no zoning and only one type is present. Twinning
on 100 is common. The extinction Z ' c is 38-41° and 2 V is somewhat variable
in different grains from 42° to 48° (measured on the universal stage) and y
— 1*720. The data indicate a pyroxene intermediate in character between
pigeonite and the diopside-hedenbergite series. In some instances, the cleav-
ages of the pyroxenes are bent and the crystals show incipient fracturing
(fig, 16A). Alteration to a fibrous greenish uralite is common, proceeding
from the periphery to the centre and often completely replacing the pyroxene.

Plagioclase is the most abundant constituent. It is in long columnar
and lath-like crystals with ^veil-defined edges in the ])rism zone and poorer
terminal faces. Fine lamellar twinning, both albite and pericline, together
with simple C’arlsbad twinning are commonly developed. The main felspar
is a labradorite aljoiit Ab 5 QAu 5 Q, but it almost invariably exhibits a gradational
normal zoning, which in some instances has been to a periphery as sodic as-
AbgoAiiQo, although the general range is from AbgQAngQ to AbypAngQ, In many
instances the plagioclase is completely replaced l3y a fine granular saussurite
and in such instances there may be a thin outer rim of sodic plagioclase. The
sassuritisation, although generally irregularly distributed throughout the
grains, seems to commence at the centre. Arborescent growths of a pale
greenish weakly birefi-ingent chlorite along cleavages and fractures of the
plagioclase are very common. The chlorite appears to come from the fibrous
uralitic coating of the pyroxene and its distribution points to the migration
of the end stage ^’apou^s and litjuids along the cleavages of the earlier crystal-
lised felspar.

Iron ores, mainly ilmenite, may occur in skeletal crystals as large as the
largest pyroxenes. In some highly uralitised types it is altered to leucoxene
(pale greyish in reflected light). Magnetite is present in one dyke to such an
extent as to influence compass readings taken in its immediate vicinity. This
magnetic feature of the dyke rocks was only noted in this one instance. It is
probable, howe\’cr. that much of the more common ilmenite has magnetite
associated with it. The iron ore when in contact with felspar is usually rimmed
with greenish fibrous chlorite. The only other ore mineral is pyrite, which is
of rare occurrence, occurring in small cubic crystals.

Qvariz varies from 0 per cent to 10 per cent. It is in angular grains
filling the interstices between plagioclase prisms. In the more acidic types
it commonly occurs in micropegmatite (along with a dusty acid plagioclase).
Angular grains of micropegmatite up to 2 or 3 mm. diameter have been ob-
served. It usually carries needle-like inclusions of apatite.

Amjyhibolc is represented by two varieties : —

(1) A pale greenish, fibrous uralite with patchy coloration developing
around the edges of pyroxene plates and frequently completely
replacing that mineral. Although the edges become frayed, the
ophitic intergrowth of plagioclase (usually saussuritised in the
uralitic rocks) with these uralite pseudomorphs still remains visible.

Geology and Petrology of Part of Toodyay District, W.A. 127

(2) A brownish well-crystallised hornblende, only found in those doler-
ites which have abundant end stage mioropegmatite. Although
at times it appears to be of primary crystallisation, it often
develops from the outer edge of the fibrous uralite, where this
is in contact with the micropegmatitic mesostasis. It has pleocliro-
ism — X yellowish-brown ; Y brownish-gi-een ; and Z bluish-green.

Biotite is a rare constituent only found in the most acid types, closelv
associated with the acid mesostasis.

Olivine is extremely rare, having been recognised in one rock only, where
it is completely replaced by a dark greenish serpentine.

Apatite is the only accessory generally developed. It is in needles except
in the most acid type, in which it builds stout euhedral prisms (fig. 16B).

Epidote, in addition to its occurrence in saussuritised plagioclase, often
occurs in narrow veinlets cutting directly across all the above-named minerals.

Chemical compositioii of the quartz dolerites.

A typical example (No. 15424) showing only a very small amount of
quartz and slight uralitisation was analysed and the result appears in Table 5,
where it is compared witli the average analysis of the Whin Sill rocks. The
analysis of the Toodyay rock closely approaclies the Whin Sill t;^’pe, differing
only in the somewhat liigher lime and magnesia in the former.

TABLE 5.

Analysis of quartz dolerite from Toodyay compared ivith the average composition of the Whin SUL

1. 2.

SiOa 49

TiOg
AlgO,
Fe^O,
FeO
MnO
MgO
CaO

NagO

K/)

H,0+

HgO-

P.O.,

FeS,

Others

1
13
3
8
0

7-64

11-84

13

27

13

65

95

15

72

16

72

04

0-14

0-45

99 • 99

50-52
2-39
76

13

3

8

0

5

9

87

•50

16

42

09

2-42

0- 96

1- 51
0-76
0-26

0-69

100-31

Norm :

Q ...

Or

Ab

An

(li ...
hy

ms

ir

ap

py

(hiartz dolerite (s[)ee. 15424), Toodyay
Average ^\'hin Sill Ty])e (Holmes and H

3-48
1 -ll
14- 15
27-52

24-88

18-33

5-34

2-43

0-.34

0-45

W.A. Analyst, K. T. Prider.
arwood. 1929, p. 539).

The resemblance of the Toodyay tpiartz dolerite to other doleritic rocks
of the Darling Range near Perth which are considered to be comagmatic, has
been noted in a in-evious ])aper (Prider, 1941, p. 46).

128

Rex T. Pkider.

(6) Ultmbasic intrusioes.-— There are two types of uitrabasi(i iiiti*tisives-
iu this area : —

(i) Completely metamorphosed types, now monomineralic tremolite-
and actinolite schists, occurring as sill-like intrusions into the
.Jimperding Series. The age of these rocks is unknown, but as will
be shown below, they are probably earlier than the later normal
granite.

(ii) Serpentine dykes. One only of tliese serpentine dykes has been
noted in the area mapped. It is about -J-chain wide and has the
same trend as the quartz dolerites. Its age relations with the-
dolerites are not known.

(i) Mctmiforphosed idtrabasic sills. -Sevcu'al narrow sill-like intrusions,
in the (piartzite were noted in the \'icinity of the R. W. Trig station. They
are jiale greenish, soft, scliistose rocks composed almost entirely of a jmle
greenish actinolite with Z A c ^ 10'^ ; y ^ 1 • 6o0 ; a = I -030 ; (--) 2V large.
This actinolite occurs in a felted network of line prisms with the interstict^s-
filled with a pale green optically positi\e chlorite. Irregular-shaped grains,
of magnetite are scatter<Hl uniformly throughout.

Another rock (15(>41), also a sill in ((uartzite, consists largely of a felted
mass of tremolitt' ])i-isms with a ground of flaky antigorite dusted with mag-
netite' inclusions.

A similar type is found just Ixdow the andalusite schist in the S.W . pai't
of tlie area and this may be the intrusive which has led to tlu' formation of
andalusite. If so, the retrogressive clianges in the andalusite, induced by
the intrusion of the normal granite fixes these ultrabasic sills as pre-granite

in age.

(ii) iSerjicntiites.- -These rocks are found in a narrow dyke Cchaiu wide
and the only a\ ailable specimen is from a ]>oiut 2S chains N., 132 chains K., of
datum.

It is a fine, even, dark greenish rock with a conchoidal fracture. Numerous
minute silvery chlorite flakes are visible in hand specimen. Under the micro-
scope it is seen to be made np of a very tine-grained aggregate of fiak>' anti-
gorite - 1*57) with occasional relict prisms of tremolite and a later develojx
ment of chlorite flakes with magnetite inclusions coating the well-marked,,
crumpled 001 cleavages.

The chlorite is a very pale greenish variety and has y — 1-595 ; «
1-589; y - a = -000: and Z 1 001. This mineral, therefore, is a slightly
aluminous iron-bearing antigorite.

The rock is a normal serpentiii'/ as shown by the following analysis ot
Specimen No. 15425.

SiCb ...

TiO^

AI.O3

FcO ...
.\liiO
CaO ...
NaoO
K2O ...

H,0-b

B,0-

MnO

f.o.

40-33
0-28
2-75
5-43
5-48
33-39
I -29
Ad
Nil
JO- 16
0-10
0- 14
Ad
0-10

Analyst : K. T. Pridcr.

99-45

(iKOUKJY AND Petrology of Part of Toodyay District, W.A. 12!^

IV. DCONOMIO DKOLOGY.

Gold and tlie refractory minerals andaliisite and sillimanite are the only
minerals of economic importance which have been noted in this area.

(1) The gold deposits occur in the vicinity of Yinni<ling Creek in the south-
west('rn corner of tlie area mapped and have be('ii described by Blatchford
(1932), Forman (1935), and Prider (1934, p. 73), and require no further descrip-
tion here. Tlie aurift^rous deposits appear to be gem^tically related to tlu'
younger non-foliated granite of the south-west part of the Area.

(2) Refractories — {a) *S'?7/iwan?Ve.- -Refi’actory clays derived from tlu'

alteration of sillimanite schists of the Jimperding Series have been workc'd
at (lackline for some years (Simpson, 1935, ]). 11 : Matheson, 1938, p. 13).

The sillimanite content of these clays is estimated to vary from 5 to 10 jx'r
C('nt of th(' rock.

In tl.(' 'Foodyay area, the metamorphism of the Jimperding Series has
been effectc'd umler sillimanite zone conditions. Although sillimanite occ‘urs
in the mir*ac('ons schists throughout tlu' area, the best sillimanite deposits-
not('d during tht' survey were in the country lying to the north-west of Key
Farm hom('st('ad, wlu're there has been a thickening of the incompetent mica-
sillimanit(' schists during folding. Although rapid variations in the silli-
manite' conte-nt are' noticeable, the average sillimanite contemt o\('r com-
paratively wide* bands is high and in certain bands u]i to 10 fea't wiele', the
sillimanite (mainly fibrolite;) forms up to 70 pov vont of the re)ck. This mate'i’ia l
constitutes a \'aluable ore' provided the small nuiscovite-biotite cejutemt can be'
se'paratc'd. I^^xperime'ntal work in this connection is at j>re>se'nt in progre'ss.

{h) Aadalvsife . — This minen-al is elevelope'd in the uppe'rmost band of
mie-a scliist in the south-we'ste'rn corner of the are'a. Tliere eloes not a])])e'ar,
in the area ma})])e'el, to be' any natural concentration of this mine'ral, suflicie'nt
to constitute* an economic proposition. Howe've'r, in the westerly exte'nsie)ii
e)f the anelalusite' mica scliists, a mile or so west of the eelge* of the mappe-d are'a,
in the* lu'adwaters of Mortigup Brook, considerable deposits of white* wc'athere'el
anelalusite schist witli abundant anelalusite are expose'el. Tlieso eleposits ai'c
be'ing e'xple>ite*d at the^ preseait tinier by a loe-al company.

V. SUMMARY AND (’0NCLUS10N8.

A e-onfe)rmable^ se'ries e>f metamorphic re>cks lias bex'n describeel. It
inclueles both sexlimemtary anel igneous type-s. The former incluele inte'rbe'elde'd
sillimanite*- anel anetalusite-mica schists, extre'inely ]iure' ejuartzite's, and
eice^asiemal e-ale^are'oeis sanelstone's, now re'pre'sentevl by e^alc-silie'ate* reie'ks.
All these rocks lie within the sillimanite zeme*. Rare biotite'-pIagie>e*lase seJiists
have* be'em noteel, in erne instance etarrying a e*onside'rable amount of ge'elrite
and cummingtemite. This biotite-ge'elrite-cummingtonite-plagioe-lase' schist is
consiele're'el te> n'sult probably from the re'gional me'tamorjihism eif a somewhat
eleilomitic clayey se'elimemt.

Interbe'eleled with the se'elime'nts there are* basic, igneeius roe^ks now
iepre>se'nttHl by schistose plagioclase amphibe>lite*s, which have be'e'u derivt'd
by re-gional me'tameirphism, uneU'r sillimanite' zone conditions, from tholeiitie:
roerks which may have be'e'ii eiriginally sills e>r flows. Thc'y are oleUr tlian thei
iirst granite intrusiem anel have bee'n folde'd along with the associateel sedimeaits.

Wide'i- bands of granite* gne'iss have^ re'sulte'd from tlie intrusion of granitic
magma into the' above rocks. The pe'rioel of intrusion coincided with tlu;
ore)ge*nic jicriod, wlu'n the pre-existing rocks wore a!te*re>el te> their pre'se'iit
state. The granite*, a porphyritic microe*line type, was intrude'el unde'r stre'ss
anel ]ire'sents a fhixiemal structure, which in the uppe'r, more narrow banel is

130

Rex T. Prider.

emphasised by the sliglit slieariiig out of the microcline phenocrysts into
inicrocline augen. Earth movements had practically ceased by the time that
the granite had finally consolidated. Late stage aplites and pegmatites were
associated with this period of intrusion. This intrusive granite has picked
up fragments of the older rocks, comprising : —

(1) Plagioclase amphibolites and related types, derived from basic

igneous rocks of doleritic composition.

(2) Altered ultrabasic igneous rocks, now represented by anthophyllite-

hypersthene and anthophyllite-cordierite rocks.

(3) Metasediments in the form of banded eulysites (derived from banded

iron ores) and biotite granulites (derived from psammitic sediments).

Tliere is no evidence of any tectonic movements of later age than the
granite gneiss intrusion.

Later these rocks were intruded by a granitic magma of similar composi-
tion to the earlier orthogneisses, together with its end stage products (aplites,
pegmatites, and quartz veins). This granite has not effected any meta-
morphism of the older rocks other than the retrogressive alteration of pre-
existing andaliisite, by vapours advancing ahead of the intrusive mass. The
intrusion of this later granite is considered to have taken place at considerable
<lepth.

A later period of igneous activity is represented by dyke intrusions of
quartz dolerite, which in their constancy of trend appear to have come up
along joints or other lines of weakness, approximately parallel to the tectonic
strike. Certain ultrabasic dykes are believed to be also post-granite.

Tlie structure of the area, which is one of abnormal strike in the Western
Australian shield, is interpreted as a major anticline, with its axis striking
a little W. of N. and pitching to the south. This structure has a recumbent
syncline on its eastern limb and is cross-folded on axes trending N.N.E.

The geological liistory of this region may be summarised thus : —

Early Pre-Cambria?i (Kal- 1.
goorlie-Yilgarn Period)

o

3.

4.

o.

Late Pre-Cambrian (Nul- (5.

lagine Period)

7.

? Aliocene .... .... S.

Present .... .... .... 9.

Deposition of argillaceous and aren-
aceous sediments with a minor
calcareous facies and with inter-
calated lava flows (or sills).

Diastrophic period accompanied by
intrusion under orogenic stress, of
a porphyritic microcline granite.
Main period of regional metamor-
phism. No orogeny after this period.

Intrusion of end stage products of the
microcline granite magma.

Intrusion of the later granite. No
further metamorphism.

Intrusion of end stage products of the
later granite. Period of formation
of auriferous deposits.

Intrusion of c|uartz dolerites.

Erosion period.

Region reduced to a p(meplain. For-
mation of laterite (Woolnough,
1930, p. 125).

Erosion periotl to the present. The
region is now a dissected, laterite-
capped })lateau.

Geology and Petrology of Part of Toodyay District, W.A. 131

VI. ACKNOWLEDGMENTS.

The mapping of the area described in tliis paper has been done by parties
of senior students in the Dept, of Geology of the University of Western Aus-
tralia during the past ten years and the contributions made by these vai*iou.s
parties are gratefully acknowledged. The greater part of the laboratory work
was done in the Dept, of Mineralogy and Petrology at Cambridge, England,
during the tenure of a Hackett Research Studentship from the University of
Western Australia, and publication of the paper has been made possible by a
grant from the Hackett Fund.

1 wish to express my thanks to Professor C. E. Tilley for assistance and
constructive criticism during the preparation of this paper, and to Dr. F. C.
Phillips for advice in connection with the petrofabric analyses of some of the
rocks examined. My thanks are also due to Professor E. de C. Clarke, who
first suggested that the Toodyay District should be closely studied, and who
has been closely associated with all the field work undertaken in this locality.
I wish to thank Professor Clarke also for his guidance in my preliminary
studies of these rocks, for his interest and encouragement in the work, and
finally, for his assistance during the revision of tlie text.

132 Rex T. Prider.

VII. LIST OF REFERENCES.

Bastin, E. S., 1909. “ Chemical composition as a criterion for identifying metamorphosed

sediments.” Joxirn. Geot, Vol. XVII., pp. 445-472.

Bayley, W. S., 1895. “ The basic massive rocks of the Lake Superior Region.” Journ,

Geology^ Vol. Ill,, pp. 1-20.

Blatchford, T., 1932. “ Report on an alluvial gold find on Yinniding Creek, eight miles

south-west of Toodyay.” Ann. RepL G.S. West. AusU for 1931, p. 9.

Broch, O. A., 1926. “ Ein Suprakrustaler Gneiskomplex auf der Halbinsel Nesodden l>ei
Oslo.” Norsk. Oeol. 'Tidskr.y Bd. IX., Hft. 2, pp. 81-219.,

Brogger, W. C., 1935. “The South Norwegian Hyperites and their Metamorphism.”
Norsk. Vidensk. Sk'r., 1934, No. 1, pp. 1-421.

(3arke, E. de C., 1930. “ The Pre-Cambrian Succession in some parts of Western Australia ”

Rept. Aust. Assoc. Adv. Sc. for 1930, pp. 155-192.

(Y»oke, H. C., James, W. F., and Mawdsley, J. B., 1931. “ The Geology and Ore Deposits

of the Rouyn-Harricanaw Region, Quebec.” G.S, Canada, Mem. No. 166.

Doelter, C., 1917. “ Handbuch der Mineralchemie.” Bd. II., 2 Hfte., Dresden & Leipzig.

Kskpla, P., 1936. “ A paragenesis of Gedrite and Cummingtonite from Isopaa in Kalvola,
Finland.” Bull. Comm. Geol, de Finlande., No. 115, pp. 475-487.

Fairbairn, H. W., 1937. “ Structural Petrology.” Queen’s University, Kingston, Ontario.

Feldtman, F. R., 1919. “The Clay Deposits at Bolgart.” G.S. West. Aust. Ann. Prog.
Rept. for 1919.

Flett, J. S., and Hill, J. B., 1912. “ The Geology of the Lizard and Meneage.” Mem.

G.S. Eng., explan, sheet 359.

I'orman, F. G., 1935. “ Lode mining at Yinniding Creek, Toodyay District.” Ann. Rept.

G.S. West. Aust. for 1934, p. 12.

Forman, F. G., 1937. “ A Contribution to our Knowledge of the Pre-Cambrian Succession

in some parts of Western Australia.” Journ. Roy. Soc. West. Aust., Vol. XXIII., pp.
xvii.-xxvii.

Goldschmidt, V. M., 1911. “Die Kontaktme.tamor})hose im Kristianiageheit.” Vidensk.
Skr., 1911, No. 1.

Hall, A. L., and du Toit, A. L., 1923. “ On the Section across the Floor of the Bushveld

Complex at the Hartebeestpoort Dam, West of Pretoria.” Trans. Geol. Soc. South Af-
rica, Vol. XXVI., pp. 69-96.

Marker, A., 1932. “ Metamorphism.” London.

Henry, N. F. M., 1935. “ Some data on the iron-rich hypersthenes.” Min. Mag., Vol.

XXIV., pp. 221-226.

Holmes, A., and Harwood, H. F., 1928. “ The Age and Composition of the Whin Sill

and the related dykes of the north of England.” Min. Mag., Vol. XXL, pp. 493-542.

Hutton, C. 0., 1940. “ Optical Properties and Chemical Composition of Two Micas from

Westland, South Island, New Zealand.” N.Z. Jour. Sc. Techn., Vol. XXL, pp. 330B-
331B.

Joplin, G. A., 1935. “ A note on the Origin of Basic Xenoliths in Plutonic Rocks, with

special reference to their Grain-size.” Qeol. Mag., Vol. LXXIL, pp. 227-234.

Matheson, R. S., 1938. “ Report on the Clackline Firebrick Clay Pits (South-West Di\ i-

sion).” Ann. Rept. G.S. West. Aust. for 1937, p. 13.

Miles, K. R., 1938. “ The Geology and Physiography of the Lower Chittering Area.”
Jour. Roy. Soc. West. Aust., Vol. XXIV., pp. 13^1.

Partridge, F. C., 1937. “ Note on the Green Micas of North-Eastern Transvaal.” Trans.

Geol. Soc. South Africa, Vol. XXXIX., pp. 457-460.

Peach, B. N., and Home, J., 1907. “ The Geological Structure of the North-West High-

lands of Scotland.” Mem. G.S. Gr. Britain.

Phillips, F. C., 1930. “ An Association of Anthophyllite and Enstatite.” Geol. Mag.,

Vol. LXVIL, pp. 513-516.

Phillips, F. C., 1937. “ A Fabric Study of some Moine Schists and Associated Rocks.”

Q.J.G.S., Vol. xciii., pp. 581-620.

Geology and Petrology of Part op Toodyay District, W.A. 133

Prider, R. T., 1934. “The Geology and Physiography of the Jimperding Area.” Journ.
Roy. Soc. West. Amt., Vol. XX., pp. 1-16.

Prider, R. T., 1940. “ Cordierite-anthophyllite rocks associated with spinel-hypersthenites

from Toodyay, Western Australia.*’ Qeol. Mag., LXXVIL, pp. 364-382.

Prider, R. T., 1941. “The Contact between the Granitic Rocks and the Cardup Series
at Armadale.” Joum. Roy. Soc. West. Aust., Vol. XXVII., pp. 27-55.

Read, H. H., 1936. “ The Stratigraphical Order of the Dalradian Rocks of the Banffshire

Coast.” Geol. Mag., Vol. LXXIII., pp. 468-476.

Richarz, S., 1927. “ Gnmerite rocks of the Lake Superior Region and their Origin.” Journ.

Geol., Vol. XXXV., pp. 690-707.

Sederholm, J. J., 1916. “ On Synantectic Minerals and related phenomena.” Bull. Comm.

Qeol. de Finlande, No. 48.

Simpson, E. S., 1926. “ Contributions to the Mineralogy of Western Australia, Ser. I.”

Journ. Roy. Soc. West. Aust., Vol. XII., pp. 57-66.

Simpson, E. S., 1928. “ Contributions to the Mineralogy of Western Australia, Ser. III.”

Journ. Roy. Soc. West. Aust., Vol. XIV., pp. 45-46.

Simpson, E. S., 1931. “ Contributions to the Mineralogy of Western Australia, Ser. VI.”

Journ. Roy. Soc. West. Aust., Vol. XVII., pp. 137-149.

Simpson, E. S., 1936. “ Contributions to the Mineralogy of Western Australia, Ser. IX.”

Joum. Roy. Soc. West. Aust., Vol. XXII., pp. 1-18.

Simpson, E. S., 1937. “ Contributions to the Mineralogy of Western Australia, Ser. X.”

Journ. Roy. Soc. West. Aust., Vol. XXllI., pp. 17-35.

Tilley, C. E., 1921. “The Granite Gneisses of the Southern Eyre Peninsula (South Aus-
tralia), and their Associated Amphibolites.” Q.J.G.S., Vol. xxvU., pp. 75-134.

Tilley, C. E., 1927. “ Vesuvianite and Grossularite as products of regional metamorphism.”

Qeol. Mag., Vol. LXIV., pp. 372-376.

Tilley, C. E., 1936. “ Eulysites and related rock types from Loch Duich, Ross-shire.”

Min. Mag., Vol. XXIV., pp. 331-342.

Van Bemelen, J. M., 1900. “ Uber das Vorkommen, die Zusammensetzung und die Bildung

von Eisenanhaufungen in und unter Mooren.” Zeits. Anorg. Chem., Bd. XXII., pp.
313-379.

Winchell, A. N., 1933. “ Elements of Optical Mineralogy, Part II.” New York.

Wiseman, J. D. H., 1934. “ The Central and South-West Highlands Epidiorites : a study

in Progressive Metamorphism.” Q.J.G.S., Vol. XC., pp. 354^17.

Woolnough, W. G., 1930. “ The Influence of Climate and Topography in the Formation

and Distribution of Products of Weathering.” Geol. Mag., Vol. LXVII., pp. 123-132.

134

Kkx T. Pridek.

Plate n.

Microcline granite gneiss.

A. Crushed microcline mosaic with sill-like quartz of
crystallisation. Nicols crossed. X 20.

B. Later quartz veinlet cutting microcline phenocrvst.
X 25.

post cataclasi.s
Nicols crossed.

Geology and Petrology of Part of I’oodyay District, W.A. 135

Plate II,

136

Rex T. Pkider,

IMate I LI.

Mierocline granite gneiss.

C. Small porpliyroclast of microelinej with granulated periphery,
crossed. X 25.

R. Finely granulated mierocline granite gneiss. Elongated uncrushed
grains abundant. Nicols crossed. X 25.

Nicois

quartz

Gkology and Petrology of Part of Toodyay District, W.A. 13?

Plate in.

JOURNAL ROYAL SOCIETY OF WESTERN AUSTRALIA

V0L.28, 1941-2. PAPER 4. PLATE I

/OOr/.

MARDIE.

PERDING

Geological Map

Toodyay Area

LEGEND

HORNBLENDE-SCHISTC
QUARTZITE C

UPPER GNEISS □

LOWER GNEISS El

BASIC LENSES E>

-E ROADS

LATERITE EZI
DOLERITE EH
GRANITE EH
MICA-SCHIST EH
TRIG STATIONS A
RAILWAY S

’ . noondeening

SECTIONS

E-W Section
E~W Section

datum

datum

A \ “V

TX — — t

\

V

\

«

f

l>i

I

Hevisiox of T!ik FhrBlOPTERA OF \Ve'=^tfrx At:rtraija.

130

5._rEVISION op the embioptera of western

AUSTRALIA.

By Consett Davis, M,Sc.

Loeturer in Biology, New England University College, Armidale, N.S.W.
Read: 9th December, 1941. Published: loth March, 1944.
Communicato'd by L. Glauert.

INTRODUCTION.

The Order Embioptera is well represented in Western Australia, somewhat
sporadic collecting having to <late nwealed five species, of which three are
endemic and rather bizarre in structure. Interesting collections have been
received from time to time from the Western Australian Museum, by courtesy
of the (’urator, Mr. L. Glauert, to whom my thanks are due. It is the purpose
of this paper to collate the systematic records relating to those species known
fi-oin Western Australia, and to add certain new locality records. It is hoped
that this may lead to further collecting, which would almost certainly result
in the discovery of new' forms.

GENEBAL ORGANISATION.

The Embioptera form a small but distinct Order of Orthopteroid insects,
showing some affinity to tlu^ Isoiitera and to a less extent the Dermaptera and
Perlaria, from all of which, how'ever, they are probably separated by a long
line of independent ancestry. No distinctly descriptive common name has
been applied to these insects ; the name “ Foot-spinners ” is here proposed,
referring to the distinctive habit of members of the Order of spinning silken
webs from the glands of the fore tarsi.

Apart from the Palaeozoic Sub-Order Pi*otembioptera, with tw'o monotypic
genera (Frotembia Till., Tillyardemhia Zal.) of ill-defined structure, the Order
is represented by about 140 Tertiary and Recent species, distributed through
some 36 genera and seven families, of which tw'O are represented in Western
Australia, each by a single genus.

The females of Tertiary and Recent genera (Sub-Order Euembioptera)
are wingless and larviform, and possess no good taxonomic characters. The
males are usually w'ingod, occasionally wingless, w'hile a few species are known
within w'hich both forms occur. The tw'o pairs of wings are equal and widely-
spaced, and the veins are rather weak. A short subcosta is present ; the
i-adius (Ri) is the strongest vein in the wing, its sector arising near the base
and forking a little before half the lengtli of the wing to an anterior branch
(Ra+d* usually fusing with Rj subterminally, and a posterior branch {R 4 + 5 ),
w'hich may be forked or, as in all Australian species, sim]->le. The media
(M) arises separately near the radial sector ; it is rarely forked, usually simple,
and often only weakly developed. The first cubital (Cui) has a strong main
axis and an anterior branch (Ciq^), usually weak, but forked once or several
times in some exotic genera ; there is no second cubital. One small anal vein
is present. Cross-veins may be frequent, but their number and position shows
individual variation, and they are of no taxonomic significance.

k 40/43

140

(’ONSKTT I)AVrS,

The tarsi of all lepjs are tlireo-st'Kinented. Tlie first sejgment of the fore
tarsi is gn^atly inflated, housing tlie spinning-glands, which discharge by
hollow setae on the plantar surface, 'fhe seconil l(*gs are wiakly developed.
The liind femora are always greatly swollen, housing the (mlargcd depressor
tibiai! muscles by whose contra(;tion the irisiud is able to dart backwards
very rapidly. The first two segments of the hind tarsus have their plantar
surfaces variously beset with vesicles and stiff setae, these being important
systematic characters.

The male terminalia, the most important taxonomic criteria, are best
understood by reference to the lar\'al form, with which the adult female
agrees essentially except in the yiresence of a transverse genital aperture
on the posterior part of the eiglith alxlominal stisnite. In tlie larva, tlie
ninth abdominal tergite is transverse, the tenth subtriangular. The ninth
abdominal sternitt; is subcpiatlrate, th(i tenth divid('d by a median longitudinal
cleft to t-wo triangular hemisternites. Bidwei'n thf'se and the bases of tlie
cerci are subannular sclerites refern'd to as cercus-basipodites. These are
very probably paraprocts, although American workers (e.*/., Snodgrass, 1035,
Fig. 140F) refer to the structures heri' intin-preted as hemisternites of the
tenth segTnent as paraproeds. Only a study of the late* embryological stages
can finally decide this point, which is, however, unimportant in the internal
classification of the Order. The cerci are two-s(igmented, each segment smooth
and subcylindrical, the basal one thicker.

In all species except Clothoda nohilis (Gerst.) (Amazon region), the tenth
abdominal tergite of the male becomes cleft at the last eedysis, although fre-
quently not to the base, so that two more or less di.stinct hemitergites are
formed ; these are furnished with cojnilatory processes. The right cercus is
usually little modified at the last eedysis, but the first segment of the left cercus
is often greatly changixl, usually b(‘c.oming clavate and often dev{‘loj)ing
echinulation on the inner surface. The second segmcxit may remain unclianged
or, as in some Australian and Nortli American genera, may be partly or wholly
resorb(.‘d into the first to form a comyjound structun'.

The ninth alxlominal sternite sends out a distal process, whicli can inde(‘d
be detected during the two previous instars in an incipient condition. Tliis
process may possibly be equivalent to fu.s('d gonocoxites, so that the whole
structure may be regarded as a hypandrium. Dor.sal to this jjrocess is the male
genital aperture, usually situated among ill-defined membraneous .structures.
The right hemisternito of the tenth abdominal segment, and the right cercus-
basipodite, usually in part degenerate, sometimes remaining as small .sclerites
unimportant functionally and taxonomically. llie left hemisternite and cercus-
basipodite may r(;main di.stinct in the adult and develop processes, but more
frecpiently they appear to form a composite structure, often with one or mor('
proces.ses, and usually referred to simply as the left cercus-ba,sipodite.

The male terminalia are in some species probably the most cojnplex in the
Class Insecta, the processes of the hemitergites, hypandrium, cerci, and left
cercus-basipodite all a,ssisting in holding the unspecialised ftunale terminalia
during copulation.

Members of the Order feed prcxlorninantly on dead vegetable matter,
more particulai-ly bark and fallen k'aves. Branching cylindrical tunnels of
silk are .spun among the food material by means of the tarsal glands, and in
these the insects live, being gregarious and subsocial, the female guarding the
eggs and young larvae in an enlargement of the gallery. Most of the Western
Australian records represent winged males, taken either at light or swarming
under weather conditions .such as induce this phenomenon in Termites. Searcli

Kevisiox of the E-Mbiofteka of Western Australia.

141

for the actual colonies may reveal wingless forms of the adult male in species
where at present the winged form alone is known, or even new species entirely
v^ingless in all stages, such as in the Eastern Australian genus Metoligotoma.

SYSTEMATIC^.

Family OLIGOTOMIDAE Enderlein 1909.

Zool. Anz., 35, p. 190. Type genus, Oligotoma Westwood, 1837, Trans. Linn.

Soc. London, Zool., 17, p. 373.

In addition to the type genus, whose diagnosis is given below, this family
includes only the genus Haploembia Verhoeff (Shores of Mediterranean and
Black Seas), which differs from Oligotoma in having the males always, instead
of exceptionally, wingless, and in the presence in both sexes of a medial bladder
or vesicle on the plantar surface of the first segment of the hind tarsus.

Genus OLIGOTOMA "Westwood 1837.

Loc. cit. (as subgenus of Embia Latreille, 1829). Raised to generic rank, Bur-
meister, 1839, Handbuch der Entomologie, Bd, 2, p, 770.

Males winged (more rarely, winged and wingless forms occur within the
same species), R4+5, M, and Cui„ simple and subobsolescent (each represented by
little more than a median row of macrotrichia in a broad band of pigment, Plate
] , Fig. 1) ; tenth abdominal tergite partly divided to hemitergites, the cleft not
extending forward to the ninth tergite ; right hemitergite with outer margin
produced back as a slender sclerotized lobe, basally overlying an inner mem-
braneous flap, the latter sclerotized only medially, this sclerotization con-
tinuous with that of the outer process. Left hemitergite with a prominent
process, sometimes complex. First segment of left cercus subcylindrical
to clavate but never echinulate ; second segment, and both segments of right
cercus, elongate-subcylindrical and distinct. Structures at the base of the
left cercus, conventionally referred to as the left cercus-basipodite, often
complex, and probably always including elements of the left half of the tenth
sternite of the larva.

Both sexes with plantar surface of first segment of hind tarsus carrying only
the terminal l)ladder, remainder of surface carrying many stiff setae (Plate
L, fig. 2).

The genus is indigenous in the warmer parts of Asia, throughout Australia,
in the islands between Asia and Australia, and islands in the Indian Ocean.
It is now tropicopolitan, spread by man.

Oligotoma glauerti Tillyard, 1923 (Plate I., figs. 5-7).

Journ. Froc. Boy. Soc. West Aust., 9, 1, p. 64. Re-described from type series,
Davis, 1936, p. 242, Plate I., figs. 6, 13, 20, 27, and 34.

Length 9.5-10.0 mm. ; head 1.4-1. 5 mm,, x 1.1 mm. ; forewing 8.2 mm,,
X 2.2 mm. ; hindwing 7.5 mm., x 2.2 mm. Colour mid-brown, eyes black,
wings with veins or their traces bordered by pale-brown bands. Head rounded
behind ; eyes subreniform ; antennae with up to 21 segments, maximum
total length 7.5 mm. ; mandibles (Plate I., fig. 7) slender, left with three
thin inwardly-directed teetli terminally and subterminally, right with two ;
inner margin of loft mandible with a median dorsi-ventrally flattened cutting
edge. Thorax, including wings and legs, normal for the genus. Terminalia
(Plato I., fig 5) with outer process of right hemitergite ending in two closely-
approximated subacute teeth ; process of left hemitergite (Plate I., fig. 6)
ending in an anchor-shaped hooklot. First segment of left cercus irregularly

142

Co’ssETv Davis.

inelanized, distally cur\'cd inwards to a tapered obtuse beak. Ninth abdominal
sternite tapered, distally smoothly truncate, margin minut(‘h' echinulate
distally and on the right. Left cercus-basipodite obtuse, curved upwards
and inwards and slightly sjiathulate. Terminalia otherwise as throughout
the genus.

? unknown.

Locality : Miily Milly Station, Murchison Ki\'er, 2b v 22, coll. L. Glauert
{holotype W.A. Museum ; paratype (J, ^Macleay ^luseum, Sydney Uni-
versity).

Oligotoma tillyardi Davis. 1936 (Plate 1., tigs. 8-10).

Proc. Linn, Soc, A’. S'. IT., 61 , 5-6, p. 241, Figs. 5, 12, 19, 26, 33.

Length 7. 4-9. 7 mm. ; head 1.3-1. 5 mm., x 1.0-1. 2 mm. ; forevving
7. 5-8. 5 mm., x 1.9-2. 0 mm. ; hindwing 6. 1-7. 2 mm., x 1.9-2. 0 mm, Colour
yellowish-brown, eyes and wings as in 0. glauerti. Head as in O. glauerti,
antennae with up to 19 segments, maximum total length 4.6 mm. Dentition
of mandibles (Plate I., tig. 10) resembling O. glauerti. Thorax including wings
and legs normal for the genus. Terminalia (Plate I., fig. 8) with posterior
process of right hemitergite as in O. glauerti ; process of left hemitergite
(Plate I., fig. 9) forcipate, right-hand lobe heavily sclerotized, curved to the left
and acute terminally, left-hand lobe more dorsal in position, flat, less heavily
sclerotized, obtuse, curved downwards and to the right terminally and ex-
cavate on the concave inner side. First segment of left cercus terminalh'
incurved to form a spatludate process. Ninth abdominal sternite tapered,
rather sharply truncate, emarginate latero-distally on the left, margin echinu-
late distally aiul on right as in preceding species. Left cercus-basipodite
slender, tapered, sub-obtuse distally, fixed to left-hand margin of ninth sternite
by membrane. Terminalia otherwise as throughout genus.

9 unknown.

Localises: Morgan’s, near Mt. Margaret, x,33 (holotype Macleay

Museum ; paratype W.A. Museum, etc.) ; Belele Station, Murchison

(4 ; Annean Station, near Nannine, Murchison (1 ^J). (Last two, new

records.)

Oligotoma approximans Davis, 1938 (Plate I., figs. 1-2, 11-13).

Proc. Linn. Soc. N.S.W., 63, 3-4, p. 252, Figs. 116-119.

6.3-6.9 mm. ; head 1.0-1. 1 mm., x 0.9 mm. ; forewing 5.8-7. 2
mm., X 1. 5-1. 7 mm. ; hindwing 4. 8-5. 9 mm., x 1.4-1. 7 mm. Colour dark
brown, eyes and wings as in 0. glauerti. Head as in O. glauerti \ antennae
with up to 18 segments, maximum total length 3.5 mm. ; mandibles (Plate
I., fig. 13) of same general form as in preceding species, somewhat stouter.
Thorax includmg wings (Plate I., fig. 1) and legs (Plate I., fig. 2, hind tarsus)
normal for the genus. Terminalia (Plato I., fig. 11) with outer process of right
hermitergite sleiider, tapered, obtuse, and slightly outcurved terminally ;
process of left hermitergite (Plate I., fig. 12) basally sinuous, tlistally slenderly
tapered, with a fiat subtriangular spine arising half-way along the left-hand
margin and diretded to the left. First segment of left cercus with a small
blunt inner lobe about one-third of length from extremity, inner margin basad
to lobe concave. Ninth abdominal sternite tapered posteriorly, distally
obliquely truncate and slightly emarginate, conca\dty filled by a projecting
membrane. Left cercus-basipodite as in O. tillyardi, Terminalia otherwise
as throughout the genus.

9 luakiiown.

Kkxisiox of the Embiopteija of Western Austraeia,

14

Locality: Ereinantle, 20~23/v/35, coli. K. K. Norris (liolotype and

paratypo (J, Macleay Musouin).

Note, — The three foro^going species ar(‘ [)robal)ly tlie most ljighly*specialis(>d
in tlie genus, agreeing well with the concept tluit greater structural cliang<‘
is to be expected in those descendants which have migrated furthest frojn
the ancestral region. They may later bo separated as two distinct sul)g(m(T’a,
one containing O. (ikiuerti, the second tlu* otlu'r two species.

Oligotoma gurneyi spinulosa l)a\'is. 193(i (Plato b, fi^s. U Iti).

Op. cit., p. 239, Fig. 3.

^ Jjcngth 8. 7-13. S mm. ; head 1.7-2. 3 mm,, x 1.4 2.0 mm. ; fort'wing
7.0-10.0 mm., x 1.6--2.5mm. ; hindwing 0.0-9. 0 mm., x 1.7-2. 5 mm. Colour
as in O. glauerti or somewhat darker. Head as in preceding species, eyes
prominent ; antennae with up to 20 segments, maximum total length 4.2 mm. ;
mandibles (Plate I., fig. 10) essentially as in the ])receding species except for
the presence of a marked excavation in the ])roximal half of the inner margin.
Thorax including wings and legs normal for the genus. Terminalia (Plate
I., fig. 14) with outer process of right hemitergite terminating in two a]>j>roxi-
mated subacute teeth, the outer one slightly incurv(xl ; process of left hemi-
tergitc (Plate I., fig. 15) slender, sinuous, with a slender acute terminal spine
directed forward and to the left. First segment of left cercus with a blunt
subterminal inward projection, the inner margin basad to this sliglitly and
smoothly concave. (This structure is distorted in tlu^ slid(5 mount of the
holotype, Plate I., fig. 14 ; for natural structure see Davis, 1936, Fig. 3, LCi.)
Ninth abdominal sternite tapered posteriorly and terminally smoothly I’ounded,
right-hand margin sinuous. Left cercus-basipodite intimately fused to the
left-hand margin of the ninth sternite, subterminally ])roduced to the left as a
sharp spine. Terminalia otherwise as throughout the genus.

9 See belowx

Localities: Morgan’s, near Mt. Margaret, x/33 (holotype Macleay

Museum ; paratype W.A. Museum, etc.) ; Lake Violet, East Murchison
District, x /27 (1 c^) ; Geraldton (1 cj) ; and the following new records : Belele
Station, Murchison (3 ; Annean Station, near Nannine, Murchison (8

in association with 1 of 0. tillyaxdi, and a female, length 14 mm., colour
dark red-brown, and therefore j^robably referable to 0. gurmeyi spimdosa
but larviform and with no apparent specific characters).

Oligotoma gurneyi Frogg. 1904, subsp. ?

A single male from Lalla Rookh Station, North-West Australia (W.A.
Museum ; cf. Davis, 1940, p. 159, Figs. 5-6) is intermediate between O, gurneyi
spinulosa, and subspecies occupying a more' easterly and northerly range
(O. gurneyi suhclavata Davis 1936, op. cit., p. 240, Fig. 4 : North Australia ;
and O. gurneyi centralis Davis 193(>, op. cit., p. 237, Fig. 2 : Central Australia),
A male from Hermannsburg, Central Australia (Davis, 1940a, p. 157, Fig. 7)
is barely distinguishable fi’om the Lalla Rookh example, so that this series
may have a wide range, and deserve subspc^cific nomenclature. The first
segment of the left cercus is only barely clavate, as in O. gw'ueyi s'ubclavata ;
the left cercus-basipodite is blunt and more or less \ipcurved, suggestive of
0. gurneyi centralis. The processes of the hemitergites, however, agree most
closely with 0. gurneyi spinulosa.

Further specimens are required, especially (1) from localities between
the range of O. gurneyi spmtdosa (supra) and the localities Lalla Rookh and

144

CoNSKTT Davis.

Herinannsbiirg ; and (2) from between tliese latter localities and the type
regions of O. gurneyi snhclavata and O. gurneyi centralis respectively. Until
some extra records arc obtained, it seems premature to name the Lalla Rookh
example subspecifically.

Family NOTOLIGOTOMIDAE Davis 1940b.

Ann. Ent. Soc. America, 33, 4, p. 681. Type genus, Notoligotoma Davis 1936,

op. cit., p. 244.

Other genera included in this family are Metoligotoma Davis (Eastern
Australia), Ptilocerernbia Fried. (Java and Sumatra), Ernbonycha Navas
(Indo-China), and the fossil Burmitembia Ckll. (Burmese Amber. ? Miocene).
The family is thus Indo-Malayan in distribution ; the feature common to all
its members is the reduction of the second segment of the left cercus in the
adult male, and its fusion, to a greater or a less degree, with the first segment.
In all genera where the character has been studied, there is a medial bladder
on the plantar surface of the hind tarsus : this is common to both sexes and
all stages.

Geims NOTOLIGOTOMA Davis 1936.

Loc. cit. Genotype, OUgotoma hardyi Friederichs 1914, Rec. W.A. Museum,

1. 3, p. 241.

Australian Notoligotomidae, males winged, or with winged and wingless
forms in the same species ; M, and Cu;ia simple, the first two well-

developed ; cross-veins relatively frequent ; tenth abdominal tergite com-
pletely divided to hemiteigites, separated basally by a trapezoidal median
sclerite. Right hemitergite with a postero-median process and an inner
process, stout and echinulate, curving forward from the anterior end of its
inner margin. Inner margin of left hemitergite produced back to a long thin
subacute process. First segment of left cercus clavate and echinulate, second
less than twice as long as its average thickness, firmly set on the outer and
distal part of the first segment and not movably articulated thereto. Left
cercus-basipodite fused to left-hand margin of hypandrium, terminally pro-
duced to a subobtuse process.

Both sexes with two bladders on plantar surface of first segment of hind
tarsus, one medial, the other terminal. (The medial bladder possibly indicates
the position of the end of one segment, the basal segment thus being formed of
two segments of an ancestral condition closely fused ; this view is supported
by the fact that the medial bladder is present in all those genera which on
other characters are regarded as primitive.)

Notologotoma hardyi (Friederichs, 1914) Davis, 1936 (Plate I., figs. 3-4, 17-19).

Op. cit., p. 245. OUgotoma hardyi Friederichs, 1914, 1 c.

^Length 8.8-11.0 mm. ; head 1.5-1. 8 mm., x 1.3-1.5mm. ; forewing
7.9-10.9 mm., x 1.9-2. 8 mm. ; liindwing 6. 4-9. 6 mm., x 1.8-2. 7 mm. Colour
rather pale brown, eyes and wings as in OUgotoma. Head, with eyes prominent,
})Osterior margin slightly incised laterally ; antennae with up to 19 segments,
maximum total length 4.3 mm. ; mandibles (Plate I., fig. 19) with dentition
as in OUgotoma. outer margin sinuous. Thorax including wings (Plate I.,
fig. 3) and legs (Plate I., fig. 4, hind tarsus) normal for the genus. Terminalia
(Plate I., fig. 17) witlx right hemitergite produced inwards and backwards to a
tapered process, weakly bifid terminally ; imier process hooked, weakly
nodulose. Process of left hemitergite (Plate 1., fig. 18) narrowest at origin,
then slightly expanded, terminally tapered and subacute. Imier margin of

Kevistox op the K^ibiopteka of Westehk Australia

145

first segment of left cercus irregular and cchinulate ; second segment short,
subconical, firmly fused to first segment. Ninth abdominal sternite with a
slender hea^dly-sclerotized finger-like process dir(?cted upwards ; left ('.ercais-
basipodite with a similar process directed back^^■ards and to the left. Ter-
minalia otherwise as throughout the gemis.

? See Davis, 1936, p. 246.

Localities : Near Perth, coll. G. H. Hardy (Friederichs’ types, no longer
extant) ; Caversham, near Perth, vi/15, coll. C. Kerruish W.A. Museum,
Macleay Museum, etc. ; infra) ; Midland, near Perth, vi /36 and vii/38 (cJcJ) ;
Cottesloe, 25/v/40 (4 ; new record).

In addition to these localities in AVest(?rn Australia, the species is known
from near Nyngan, N.S.W., and from near Townsville and Rockhampton, Q.
(Davis, 1936, p. 246, and 1940a, p, 158).

Note . — It is clear that Friederichs’ types have been lost. A male (slide
mount ; Macleay Miiseum) has accordingly been designated neotype. The
locality, Caversham, is as close to the original type locality as can be de-
termined ; the specimen agrees with Friederichs’ description in all respects.

KEY TO THE SPECIES OF WESTERN AUSTRALIAN EMBIOPTERA.

(Characters arc for the adult male ; tlie tarsal features apply to both sexes and all
stages.)

1 . R.j +5 and M well-defined ; first segment of hind tarsus with a medial bladder on the

plantar surface ; first segment of left cercus echinulate, second segment of length
less than twice average breadth, not distinctly articulated with first Nofolkfotorm

hardy i (Fried).

R4+5 and M subobsolescent ; medial l)ladder of first segment of hind tarsus absent ;
first segment of left cercus not echinulate ; second segment with length at least three
times maximum breadth, and clearly and movably articulated with first segment 2

2. Process of left hemitergite with a terminal hook ... ... ... ... ... 3

Process of left hemitergite with a lateral process ... ... ... ... ... 5

3. Terminal hook of process of left hemitergite projecting to both sides from point of

attachment ; margin of ninth abdominal sternite minutely echinulate near right-

hand distal angle Oligoioma glauerti Tillyard.

Terminal hook of process of left hemitergite projecting only to the left from its point of
attachment ; ninth abdominal sternite smooth ... ... ... ... ... 4

4. First segment of left cercus clavate ; left cercus-basipodite spinescent Oligotoma

gur7ieyi spinulosa Davis.

First segment of left cercus subcylindrical ; left cercus-basipodite blunt Oligotoma

gurneyi Frogg., subsp. ?

5. Lateral lobe of process of left hemitergite blunt, spathulate ; outer process of right

hemitergite terminally bidentate ; first segment of left cercus very markedly incurved
terminally ... ... ... ••• Oligotoma tillyardi Davis.

Lateral lobe of process of left hemitergite a sharp, fiat spine ; outer process of right
hemitergite terminally tapered, obtuse ; first segment of left cercus only slightly
incurved ... ... ... ... ... ... Oligotoma approximans Davis.

jVoie. — The exotic species Oligotoma saandersii Westwood (native of the Indian Region)
and O. humbertiana (Sauss.) Davis (native of Ceylon) may later be found to occur in North-
West Australia. The terminalia of each (cf. e.g., Davis, 1939, Figs. 3, 5) are very distinc-
tive ; the former has a curved spine arising subterminally from the left-hand margin of
the ninth abdominal sternite, and curving under this and upwards on the right ; the latter
has a prominent tooth on the outer margin of the outer process of the right hemitergite,
this tooth being situated well forward of the termination of the main process.

14 ()

CoNSETT Davis.

REFERENCES.

IJiirmcister. A., 1S39: Kandbnc-h der Entomologic, Bfl. 2. Berlin.

Davis. —

1930: Studies in Australian Embioptera 1. Proc. Linn. Hoc. A\.S', B'., 61, 5-0.

I93S : Studies in Australian Embioptera, Til. Ibid., 63, 3-4.

1939: Taxonomic Notes on the Order Embioptera, I. Ibid.. 64, 1-2.

1940a : Studies in Australian Embioptera, IV. Ibid., 65, 1-2.

1940b : Family Classification of the Order Embioptera. Ann. fJnt. Hoc. America, 33, 4.
Enderlein, G., 1909 : Die Klassifikation der Embiiden, nebst morphologisehen und physio-
logischen Bemerkungen, besonder.s iiber das Spinnen dcrselben. Zool. Anz., 35."
Friederichs, Iv., 1914 : A New Species of Embiid from Western Australia. Records W.A.
Mnsemn, 1, 3.

Froggatt, W. W., 1904 : Notes on Neuroptera and Descriptions of New Species. Proc.

Linn. Hoc. A^./SMT., 29.

Snodgrass, R. E., 1935 ; Priiunples of Insect Morphology. McGraw Hill.

Tillyard. R. J., 1923 : The Embioptera or Web-spinners of Western Australia. Journ.

Proc. Roy. Hoc. West. Aust., 9, 1.

Westwood, J. O., 1837 : C3iaract‘'rs of Rmbia a genus allied to the White Ants. Trans.
Linn. Hoc. London, Zoot., 17.

EXPLANATION OF PLATE I.

Figs. 1-2., Oligotoma approximans Davis, holotype 1, right fore and hindwing,

X 9. 2, hind tarsus viewed laterally, x 38.

Figs. 3-4., Notoligotoma hardyi (Fried.), neotype ^ : 3. right forewing, x 0. 4, hind

tarsus viewed laterally, x 38.

Figs. 5-7., Oligotoma glanerti Till., paratype ^ : 5, terminalia from above, x 9. 6,

proc-ess of left hemitergite of tenth abdominal segment, from above, x 38. 7. mandibles

from above, x 60.

Figs. 8-10., Oligotoma tillyardi Davis, holotype Corresponding structures and

magnifications to Figs. 5-7.

Figs. 11-13., Oligotoma approximans Davis, holotype : Corresponding structures and
magnifications to Figs. 5-7.

Figs. 14-16., Oligotoma gurneyi spiniilo<a Davis, holotype : Corresponding structures
and magnifications to Pdgs. 5-7. (First segnunit of left cereus slightly distorted on slide
mount. Fig. 14. LCJ.

Figs. 17-19., Notoligotoma hardyi (P’ried.). neotype ^ : Corresponding structure.s and
magnifications to P"igs. 5-7.

(All figures prepared with a projection drawing appai-atus. oi- baserl on camera lucida
outlines. Setae omitted except in P5gs. 2 and 4.)

9, ninth abdominal t^rgite ; lOL, lOR, left and right hemitergites of tenth abdominal
segment ; L(\, LCg’ RG.' second segments of left and right cerci respectively).

Revision of the Embioptera of Western Australia.

147

PLATE T.

By Authority: Kobkrt H. Miij.kr, Government Printer, Perth.

•>.

. . . 1 .

t

. i

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. :S'.

C

!

Ecological Succession Obseiived Duking IIegeneraTioN
OF Triodia Pungens K.Br. After Burning.

140

6.— ECOLOGICAL SUCCESSION OBSERVED DURING
REGENERATION OF TRIODIA PUNGENS R.Br.

AFTER BURNING.

By Nancy T. Burbidge, B.Sc- (Hons.).

Head 10th March, 1942 : Published Oth December, 1943.

Certain investigations, into the sheep carrying capacity of the spinifex
country of the north-west of Western Australia, are being carried out by
ofhcers of the Institute of Agriculture of the University of Western Aus-
tralia, at Warralong Station, about sixty miles east-south-east of Port Hed-
land. The name spinifex is used locally for various species of Triodia R.Br.
In the course of a taxonomic study of this genus the writer visited the country
between Port Hedland and Marble Bar during the months May, June and
July, 1941. Some weeks were spent at Warralong. An area of 400 acres,
on this property, is being used for certain feeding experiments. The spinifex
cover has been removed by burning and the plots therefore provide an ex-
cellent ai*ea for a study of regeneration. Two quadrats (20 x 20 links) were
mapped in the experimental area and two (50 x 50 links) in adjacent country
which provided examples of more advanced regrowth.

The climate is semi-arid with high summer temperatures. Rain may
fall at any time between December and March. The fails may be spread over
some months or there may be heavy storms (willy-willies) when the average
for the year may be exceeded during a twenty-four hour period. The yearly
aggregates are, however, not notably variable and the climate may be de-
scribed as one in which dry summers are common, but prolonged droughts
rare. In 1941 a dry summer w’as followed, in early March, by a violent storm
which caused record floods in many of the rivers.

The Warralong experimental plots are situated on the granitic plain to
the south of the De Grey River. Except for an occasional ridge of stony
hills due to intrusive rocks the general landscape is flat or very gently un-
dulating. The soil is a light sandy loam, reddish in colour (Teakle, 1938).

The vegetation has been described as semi-desert savannah (Teakle l.c.).
Spinifex {Triodia spp.) is the dominant genus. T. pungens R.Br. is the
most important species and is frequently the only perennial grass present.
Small trees and shrubs are present; their relative density varies and appeai^s
to be dependent on some soil factor which is, as yet, unrecognised.

T. pungens is the only species of the genus with real pastoral value. It
is a coarse, tussock-forming, resinous grass very variable in its growth habit.
At the Warralong plots the general form is a pyramidal tussock varying in
size up to two metres in diameter and about the same in height.

The use of fire to improve pasture is, of course, practised in many parts
of the world. Old fibrous and unpalatable material is bm*nt and young re-
growth is more easily reached b}^ the grazing animal. In burning spinifex it
has been assumed, and indeed widely claimed, that the fire burns the tussock
back to a central butt from Avhich young shoots develop. While this is ad-
mittedly true in some places, the writer’s observations showed that it is by
no means ahvays so. Over a wide area fire causes the complete destruction
of the plants and regeneration takes place by means of seedling establisii-
ment.

k 41/43

150

NaNX’Y T. Bl’RBTDfiK.

Observations showed that along the coastal zone, from Port Hedland
to the northern end of the 80-Mile Beach, burning left the tussocks with a
viable butt Avhose regrowth Avas aA^ailable to the grazing animal Avithin a
feAV months. HoAvever, through the inland country along the Be Grey,
ShaAV and Coongan Rivers up to Marble Bar the tussocks are completely
destroyed and it is at least a year before regroAA'th is available to the sheep.
Under such conditions the risk of soil erosion is great. Fortunately the
Triodia tussocks are fairly AA’idely spaced (Fig. 4) and it is difficult to keep a
running fire going. Thus only small patches from tAvo or three tussocks
up to about an acre are burnt in each place. The spacing of the ground
cover has probably played an important role in protecting the vegetation
from the consequences of biotic activities during the seventy years of pas-
toral occupation.

The general custom is to burn during the mustering period in April
and June when the men, riding the paddocks, drop lighted matches at ran-
dom. Sporadic burning is carried out at any time during the year.

It is evident that, if the tussocks are completely destroyed in May or
June, the ground Avill lie bare until the following January or March Avhen
the summer rains bring about seedling germination. It aA'US hoped that a
study of the progTess of regeneration might shoAv w^hether any undesirable
changes are likely to occur as a result of the burning process. That the
vegetation may be altered Avill be shoAvn in the discussion on succession.
The alteration is a result of a combination of factors of AAhich fire is one.

Figure 1.

Ecological 8ucckssion Obslrvkd During Regeneration

OK TrIODJA PUN(iKXS H.Bk. AfTKH HURXIN(i.

lol

QUADRATS.

Eig'ures 1 and 2 (20 \ 20 links ==4x4 metres), represent the pri-
mary stage in regeneration. The Triodia seedling's are associated with
numerous annual species. The areas mapped were on the Warraldng ex-
perimental }>lots and the original co\'er was burnt oft' in December, 1940.
Rain fell in March (.willy-willy) and in April -and May (light falls). The
quadrats were ma])pGd in June. It is evident that germination took place
at more than one stage. The seedlings varied from 2-6 cm. in height.

The sandy loam had a scanty cover of drift sand, formerly associated
with tussocks. This drift sand was disturbed by the March floods. It is
probable that many seeds, imbedded in the sand, were washed away.
Whether this had much effect on the relative abundance of the various
annual species cannot be ascertained from one season's observations. The
sand drifts do, however, explain the grouping of seedlings which is evident
on the maps.

In Figure 3 (50 x 50 links = 10 x 10 metres) the patch of ground re-
liresented was about a hundred yards from the experimental plots. The
quadrat was surveyed in June. At this stage of regrowth there was fierce
competition between the Triodia tussocks and ephemeral growth Avas meagre.
In one corner an adult tussock had survived. Across another there is a
patch of primary regroAA’th Avhich suggests that it Avas burnt in the previous
season.

152

Nancy T. Burbidge

It was at least two and possibly three years since the plants on the
]-est of the (]uadrat had germinated. They were about 30-40 cm. high.

In Fig-ure 4 (50 x 50 links = 10 x 10 metres) the adult tussock stage
is represented. TriodUi pungens had estal)lished a closed community and
ephemeral growth was negligible.

The quadrat was surveyed on ground about two hundred yards from
the experimental plots. All ephemeral growth Avas dead Avhen the mapping
was done in June. Observations suggest that it was at least live years
since this patch of ground had been burnt and probably much longer. The
])lants were about 70-100 cm. high.

Country surrounding the (piadrat carried occasional plants of blood-
wood {Eucalyptus dichromophloia) , kangi {Acacia pyrijoha) and geina
( Carissa lanceolata ) .

HECIFNEEATION.

The germination of Triodia seeds is rapid after rain has suliieiently
moistened the soil. The process seems to be i elated to soil moisture and not
to any particular period of the year. However only those ]dants which
germinate early in the year form a root system extensive enough to support
it during the drought conditions of the ensuing summer months.

Ecological Succession Observed During Regenerations
OK Tkioota Pungexs K.Br. Akti:k P>urxix(;.

153

The Trioclia seedlings are soon followed by small annuals such as Mol-
higo mollnginh, Euphorbia australis, Portulava oleracea, Bulbostylis bar-
hata, Eriachne pulchella, Aristida arcnaria, Sporobolus australasicus, Era-
grostis Dielsii and Ichnanthns australiensis. Amongst perennials which are
present at this stage the most important is Cassia notabilis. This species is
an indicator of recent burning (within two or three years). As the plants
are destroyed by tei’mites they do not appear in older patches of spinifex.
Tt is possible that fire improves the germination rate of this species. Other
perennials which occur as occasional plants are Solanum diversifolium, Sida
platycalyx, Corchoriis elachoearpus, Hybanthus enneaspermns, Eragrostis
eriopoda, Eriachne obtusa^ Chrysopogon pallidus and Neurachne Clementii.

The areas chosen for the first two quadrats represented good regenera-
tion. Other ground nearby varied from heavier regrowth to none at all.
Regrowth appeared to be related to the water content of the soil Areas
burnt before or shortly after the March hoods carried good regrowth. Those
burnt later showed fewer and fewer seedlings as burning continued
through the months. Those burnt in May showed no regrowth at all. There
was no rain during June or July.

Apart from regeneration on burnt areas, regrowth was also to be seen
on claypans wherever sand di'ifts formed. As a result, these bare areas

154

NaXOV T. BURIUlKiE

usiuiUy had an irregular formation due to patelu'S of regi’owth, in various
stages of regeneration, here and there on their surface.

During the summer months following the lirst period of seedling de-
velopment soil moisture continues to play a leading part and presumably
becomes a severely limiting factor. Those seedlings whose roots have not
penetrated more than the top layers of soil soon die. It is a feature of
young Tviodiu seedlings that they have the most feeble attachment to the
soil. It is not unusual to see a tussock 10 or 15 cm. high and about the
same in diameter with only a single root holding it to the soil.

The ephemeral growth dies otf very early, the ma,]ority having finished
seeding by September which marks the commencement of the hot weather.
The perennials, including T. pungens, pass into a practically dormant state.

In the second season the young tussocks compete not only Avith each
other, but also Avith another crop of annuals resulting from the summer
rains. These include Polanisia icosand)'(x ^ Avistida arcwuriu, D act plot aenium
radnlans, Mollugo 7nolluginis, Indigofera viscosa and other species present
during the first season.

By the third season the competition between the tussocks has become
the dominant feature and annuals pluy a minor role. (Dig. d).

As the tussocks become older many die out. Others coalesce to form
compound- groups. Owing to the formation of adventitious roots at many
nodes, both on the ground and throughout tiie dump of culms, the plants
after the first year, become attached to the soil at many points Avithin their
circumference. The plants mapped in Digs. 5 and 4 all exhibited one or both
of these traits.

ECOLOGICAL SUCCESSION.

The areas denuded of plant cover as a lesult of burning represent
‘^secondary” bare areas, ecologically speaking, as opposed to ‘^primary”
bare areas such as those left by landslips or earthquakes.

In this case it is not essential that plants immigrate from other areas
to form a neAV population since the soil may already contain the elements
necessary for regeneration. This is so in the spinifex country since regrowth
comes from seeds in the soil.

The regenerative sere may be divided into the following stages: (i)

primary stage Avith annuals and Tnodia seedlings in open competition.
This coA'ers the period from germination until the rains of the folloAving
summer; (ii) secondary stage lasting more than one season, during Avhich
period the Triodia seedlings assume a dominant role and eliminate the
annuals fi’om the association; (iii) adult tussock stage Avhere the Triodia
plants have formed a closed community in Avhich there is very little or no
ephemeral growth; (iv) climax association AAdiere small trees and shrubs
(which are present in a young state in (ii) and (hi)) such as Eucalyptus
dichromopliloia, Acacia pyrifoUa, Afalaya lieyniglauca, Bolichandrone lietero-
f)hylla, Hakea lorea (as examples of the former) ; and Carissa lanceolata,
CassiU' venustUy Cassia oUgopliylla and Acacia trauslucciis (as examples of
the latter) ; are associated AAuth adult tussocks.

The effect of man and his grazing animals on the association appears, so
far, to have been very slight in comparison Avith the effect on native vegeta-
tion in other parts of the State. Nevertheless there are places, both on

Ecological iSuccKssioN Observed During KegeneraTton

OF Tkiodia Puxgens H.Br. After Burning, 15o

Warrajong- and on adjoining- ]>i’o|)crli(‘s, whore major changes in the vege-
tation can he recognised. The Triodia on these -areas has l)een destroyed
aiifl re])laeed with an Erafjvosfi.'^ eripoda-douunixni association. This dis-
turbance of tile normal se(]uence is not- merely a retrogressive change in the
succession and it seems best to define it as a di'iclim(f r. That the disclimax
is a “permanent” change is indicated l)y the fact that in one place the coarse
Avoolly bases so chai’acteristic of Eracjrosii-^ eripoda averaged 20 cm. in
diameter. A<'cording to report this ret)i'('sented more than fifteen v(‘ars of
growth. The basic causes of the change seem to lx* burning followe<l by
prolonged and h(‘avy stocking. Whether burning was a i>rimary cause in
all cases could not be proved. In this Kraprostis association seedlings of
Triodia puugevi^ were absent and it is doubtful if they could establish them-
selves against the ti(‘rce competition. That the country formerly ('arried
Triodia pnngem^ is known from rcj^ort and from the e.xistence of isolated
tussocks of this sp(‘cies among the Krarp-ostis; plants. Other jdants ])resent,
both ephemeral and perennial, were the same as those in a normal Triodia
p H nge n .9- a sso c i a t i o n .

In one ai'i'a, on the adjoining propei’ty of Kginbah, there were indica-
tions that the Eragrostifi cf/;;ode-association was being removed in its turn
by the grazing animal. The result was a growth of annuals in which MoUugo
mollaginis, Aristida aremtria, A. Inpjroiactrira and Polaniria icosandra were
the most ini])ortant species. This repi'esents a retui’ii to the condition seen
in the early stages of the sere on a bai’e area.

SUMMARY.

Burning of Triodia pangens results in either the destruction of the
tussock with regeneration from seedlings or a viable butt remains whiidi
spi’outs again almost immediately.

In the coastal region, l)etween Port lledland and the northern end of
the 80-Mile Beach, a viable butt is left, but inland along the Shaw, Coongan,
and I)e Grey Rivers the tussocks are usually destroyed.

Quadrats showing stages in i-egeneration after burning are figured and
discussed. These were mai)[)ed in or u(*ar the experimental plots at Warra-
long Station.

A s(‘re is delined. The climax is one in which small trc(‘s and shrubs
are associated with the dominant Triodia pnnfjavs. Ephemeral gi’owth is
scanty in the mature stage.

A permanent change in tin* vegetation, diu' to tire and the grazing
animals, is des{'rib(*d.

ACKNtnV LE I )G MENTS.

The work desci'ibeil in this ])aj)er was mad(‘ j^ossible by the granting
of the use of certain facilities by Professor G. A. (hxrrie and Mi*. A. M.
Stewart of the Institute of Agricultui'e, University of Western Australia.
The author also wishes to ex]>ress hex- ap])reciation of the hospitality of Mr.
and Mrs. Eraxd<. Ilardie of Warralong Station, also for assistanct' in the
Held from Mi’. Frank i\rel\ille of the Institute of Agriculture,

Nanc\' T. Burbiik k.

156

REFERENCE.

1938j Teakle, L. J. H. : A. Regional Classification of the Soils of West-
ern Australia. Jour. Roy. Soc., W.A., Vol. 24, 1937-38.

KEY TO SYMBOLS USED IN FIGURES.

Trioclia pungens seedling

. . 0

„ „ tussock

o

Sporobolus australasieus

. A

Eriachne pulchella

a

Ichuanthus australiensis

0

Eragrostis eriopoda

Bulbostylis barbata

Mollugo molluginis

.. K

Portulaea oleracea

.. /a

Polanisia icosaiulra

c

( 'assia iiotabilis

M

Cliaiitlius Dampieri

(3

Euphorbia australis

SI

Goodenia

pT

yj

•• X

Miscellaneous

.. '9

»

Ecological Notes on the Vegetation of 80-Mile Beach. 157

7._EC0L0(IICAL

NOTES ON THE VEGETATION
80 -MTLE BEAC^H.

OF

Hy Xancv T. BuRHiiKiK, I>.S(*. (Hons.).

Read lOlh March, 1941 : Published 2olh February, 1944.

The area with which this pai)er is conceiaied is the portion ol the coast
oT Western Australia lietween (^ape I\eraudr(“n, where the Xo. 1 Rabbit
Proof Fence reaches the coast and Ca])e Mississey, which marks the northern
boundary of Anna Plains Station, and is the first break in the coastline of
the i)each. The ecolo,i»ical notes were obtained during a series of eollectiiig
trips through the area during July, 1941. They cover the country to a
depth of from ten to fifteen miles. On an accomi)anying map the junction
between the two main soil types is marked, since it is also tlie junction
between the main ecological /ones. The country north of Anna Plains has
not been s(‘(*n by tlu' author. The vegetation ol the i>ortion of i^ardoo,
south of the Rabbit Proof Fence, is complicated by creeks inland and by
niangTO\(' swani})s on the coast. Fcologically it is a transition region hdween
the HO- .Mile Beach and the country along the l)e (ii’cy River, Avhich is d(‘alt
with elsewhere.

The, climatic conditions are semi-arid. The annual iirecijdlation vari(‘s
b(4ween twelve and foinleen inches for the area. Most of the rain tails dur-
ing the period December to March and , owing to the porous nature of the
soil the water soon disai>pears. Permanent water holes, excej)t for a few
small native soaks, are absent. There is a marked winter drought.

PllVSrOGRAPlIV AND BOILS.

The b('ach faces north-west or north-north-west in a long slow curve.
It is without important features throughout its length. At Wooroo Creek
there is a sharp l)ut small indentation which includes a small mangrove s.wamp.
Elsewhere the beach is unbroken. Wooroo (’reek extends inland for about
a mile. Behind the beach are series of sandhills varying from one to three
miles in depth. The first row of the series are of normal dune ty])e, both
with and without vegetation. The sand here is very like that (>f the beach.
Inland are sandhills carrying a different form of vegetation. The soil of
these contains more organic matei-ial and is a light grey in colour compared
to the creamy white beach sand. Some of the ridges have outcrops of a
sandy limestone which, from the nature of its shell content, is of very recent
character. Through the section included in Pardoo, Wallal, and Xalgi the
sandhills dominate the coastal ]dain. Intermixed with them, howevei', are
wide Hats with a light grey loam soil. Through the Xalgi area these gradually
increase in relative importance until in Anna Plains, where the coastal ]>lain
widens tmt, the loam flats are the main feature of the landscape.

The junction between the coastal ]Aain and the si>inifex pindan is marked
on the ma]). It is a A-ery abrupt chang(‘ from the grey soils of the plain
into the i*ed desert sand. fhe vf'getation also changes abi’U]Al\ and the
transitional phase occupies only a few yards. The st)inifex idndan <'ountry
is undulating or more or less Hat with, in some places, long narrow sandhills
running more or less ])arallel with each other and a varying distance apart.
The only rock seen outcropping Avas a dark red ferruginous sandstone.
Through Pardoo and Wallal the outcro))s form small hills but on Anna Plains
they result in piles of stones a few feet above the surface.

158

Ni^NCY T, BuRBIDGPi.

Across Nalgi and the southern section of Anna Plains is a curious tall
shrub zone. The zone varies from under a mile to several miles in depth.
The zone has two features worthy of comment. Firstly, just south of the
Nalgi boundary is an area with numerous “blowholes'^ from a few inches to
several feet in diameter. The depth of these is unknown. From surface
indications the limestone here appeal’s to be different in nature from that
of the coastal sandhills. Secondly, through the blowhole country there are
a. few short drainage channels. Apart from these the whole area dealt with
in this paper is entirely lacking in defined water courses.

Small ])atches of this shndiland occur in some jdaces on Wallal but
there is not a continuous zone as there is further north. There is no other
outcrop of blowhole limestone.

Other features of interest on the coastal jdain are occasional red sand
ridges. The soil appears to be intermediate in character between the pindan
sand and the coastal type. The former has possibly been left there as a
result of wind action. Ea.st or south-east winds are of almost daily occurrence
over many months of the year. Such ridges are usually near the junction of
the pindan and the plain. In some cases the ridges have outcrops of the
coastal sandy limestone.

p]COLOGY.

As has already been said the area may be divided into two main zones,
i.e., the coastal plain and the spinifex pindan. However, the whole region
belongs in the great ecological region Avhich .stretches from the Ashburton
River to the Fitzroy River. This area is nearly all spinifex country, i.e.,
species of Triodia R.Br. dominate the landscape. Along the SO-Mile Beach
there are just the two phases of the ecological type to discuss. Triodia
jningens R.Br. is the dominant species and often th(‘ only grass present.
Various other arasses are important in different sections but thev will be

Ecological Notes on the Vegetation of 80-Mile Beach. 159

discussed under their I'espective headings. The trees present are stunted and
often contorted in ai>pearance. Among the shrubs various species of the
Malvaceae are conspicuous. The Leguminoseae, particularly in the ])indaip
provide a number of shrubs and small trees. These latter are mostly spe(*ies
of Acacia.

The following spectra gi\e a picture for the area and foi’ the two zones.
It will be seen that tho’e is a large number of annual species. This is rather
misleading since the (luantity of each of these is small and some species
were only obsei’ved once, i.e., at the time of collecting. iVnother feature is
that nearly all s[>ecies are confined to one or other of the zones.

No. of
Species.

M.

N.

Ch.

11.

Th.

E.

Whole Area . .

192

15

28

9

S

37

1

Coastal Plain

65

11

12

17

9

49

—

Spinifex Pindan

181

16

34

5

8.5

34

1

A. Coastal Plain.

(a) Coastal Sandhills —

(i) Beach dunes.

(ii) Triodia sandhills.

(b) Loam Flats —

(i) Grass plains.

(ii) Samphire fiats and claypans.

(c) Cadgibut Shrubland.

(a) Coastal Sandhills.

(i) The beach sandhills are of the normal sand dune type, similar to
those seen elsewhere along the west coast of Western Australia. Those with
plant growth have a covering of Spinifex longifolms. Other plants include
Ipomaea hiloha, Euphorbia mgrtoides, and PtUotus villosiflorus. This asso-
ciation, which is often almost pure Spinifex, seldom occupies more than the
hrst row of the^ series of sandhills along the coast.

(ii) Included under this heading are all the other sandhills of the coastal
area. They include .stationary sandhills of dune origin as well as those whose
inner core of limestone is api>arent in outcrops. As has been said above these
sandhills are the main feature of the ]dain throughout Pardoo, Wallal and
Nalgi. The vegetation is dominated by tlu' coastal form of Triodia pungent^
which has long wiry leaves and a dense tussock habit. Near the coast there
is a reed like grass, Panicnm sp., mixed with the Triodia. This is less
appareiit further inland. There are* also patches of Acacia salicina forming
small thickets in the hollows between the sandhills.

On the ridges, where red .sand has been deposited, various plants are
present which, normally, are restricted to the spinifex-pindan country.
These include Baahinia C unninghamil, Acacia holosericea, Acacia transluccns
and the form of Triodia pungens which is found in the desert. This last
has shorter leaves and long trailing culms which are more resinous than
those of the coastal form.

The general picture of the coastal plain is that of undulating sandhills
with a covering of coarse tussock grasses and with small Acacia thickets in
.some of the hollows.

Xancy T. "Burbidge.

1 60

1. Coastal Plain slio^ving tlie sharp junction ‘with the
Cadgibiit ISiirubland, A'aigi.

(I>) Loam Flats.

(i) The grass plains are relati\ely uiiiin])ortant until the Anna Plains
couiitvy is i‘eaeh(Hl. Flsewhere only odd patches occur between the sandhills.
Also over a good deal of the area the native grasses have been replaced by
the introduc(‘(l speci(‘s Ceyiclirun clJiaris (Huffel (trass). i\Iost of the indcs
concerning tin* native grass(‘s wcu'e ma<le at Anna Plains Avhere the Mats have
])een grazed by catth* for some y(‘ars. Whether there has been any Moristic
alteration as a I’esnlt of the grazing it is not ))ossib](‘ to say on tin' availabh'
information.

Triodia inmytns is lacking from tin' loam soils. Tlu' grassland is a
mi.xture of species and it would be ditlicult to say which is the most important.
Species which do occur are h'ragrosfis Dirfsd', K. larunaria, Xerochloa harhata,
Enneapogon phmifoUus, EporoJxjhis actinocladus, Triraphis mollis, Dicliini-
thium humilius, Chloris rnderalis, Vanicum decompositum var. scaherriminn
and also Sporobolus virginictt.'^ which has two growth forms. As Salt Grass
it foi’ms a coarse mat 8-12 inches thick of erect culms whi^e as Coastal ('’o’U'h
grass it has long trailing culms oft('ii more than twelve feet in length. The
two forms app(*ar to be tin* i‘(*snlt of a r(‘sponse to a slight alteration in soil
piineral salt content.

EooLOGiCAL Notes on the Vegetation of 80-Mile Beach. 161

On wluit Mi)pear to be damper patches in the grassland there are colonies
of EuUdia fulva oi’ Botliriochloa dccipiens (both knoAvn as Bundle Bundle
Brass) Avith Paniciim decompositum as a subordinant form.

As a j'ule the grass flats are treeless areas but occasionally there arc
small grou[)s of Melaleuca leuofiidendron and M. latiiandra (Cadgibut). Herej
as a ruhg most ot' the grasses are absent, though Cenciirus ciliaris may occur.
These tree groui)s i'e]>resent isolated [)atches of tlu* Cadgibut shrubland facies
which forms a definite phase in the transitional zone betAveen the coastal
plain \egetation and that of the si)inifex pindan. This border type appears
not only where the Junction is betAveen loam and red sand but also Avhoi’e it
is between grev and red sand.

(ii) Samphire flats and clayj>ans occur in the sand hill country as well
as on the grass flats. The soil is grey loam. Since there is no pi'oper drainage
system the loam flats are the only places AA'here one Avould expect Avater to
lie after I’ain. Tracks through these Hats have a sinister reputation for their
mud during the Avet season. It seems likely that the soil here has a slightly
higher concentration of mineral salts. If this is so it explains the zonation
which can be observed. There are three zones. In the outer one Sporoholus
virgin'eus (Salt Grass form) is dominant and there may be colonies of
Triantliemii turgidi folia. Other }>lants Avhich have been collected in this zone
include Tricliinium exaltatnm and Scaevola spinescens. In the next zone
Trianthema turgidifolia has beconu' dominant. !Mixed Avith it are Bassia
astrocarpa and Atrijdejc elacliopliylla. In some i)laces either of; these zones
may carry small thickets of Acaoia hivenosa or Acacia salicina^ but othei’Avise
trees and shrubs are completely lacking on the samphire flats. Trianthe7na
turgidifolia is locally known as yelloAv samphire. The third zone sees
Trianthema turgidifolia replaced by Artlirocnemim Bentlumiii (Red or Black
Samphire). This species gradually thins out until the bare ground of the
centre is reached.

A good example of samphire Hat is seen at Wooroo Creek between the
sandhills. Tlu‘ portion near the creek, Avhich has a border of mangroves, is
Hooded at certain times of the year Avhen there are exceptionally high tides.
Over this portion Trianlhc'iva is replaced by Arthrocnenium Benthamii.

(c) Cadgibut Shrul)land.

This s(‘ction represents a peetdiar deA'elopment of the coastal plain.
Various sp('cies of Melaleuca are abundant and dominat<* the vegetation.
This is in sti’iking contrast to the almost treeless plain. The soil is gi'ey
loam as on tin* flats. The s])ecies of Mrhdeucff present are M. leucadendrou .
M. also})liila and M. ]asi(cu<lra. This association is found along the Nalgi
frontage betAveen flu‘ ]dain and the si>inifex i>indan. In tlu^ southern section
of Anna Plains if turns (*astAvai*ds and is lost in the desert. On Wallal isolated
patches of similar <M)untry are found in a similar situation, but there is no
distinct sub-zone.

Apart from the Melaleuca tre(‘s (up to 15 feet high) the vegetation is
v('ry much the same as on the samphire tlats. HoAvever occasional patches of
Triodia pungens of the plains form are to be seen, especially Avhere small
sand drifts have foi'ined under the tv(‘es. The local impression Avas that
Triafithema turgidifolia had replaced Sporoholu:^ virginiem^ during the graz-
ing of the yi'ars of occupation. lIoAA’over, since tin; former can Avithstand

162

Nancy T. Burbidge.

higluM' coiu'eiitrntions of mineral yaltSj one must hesitate befon' placing all
the responsibility of such a change on to the biotic factor. It is e(iually
possible that cliniati(' cycles may be an important factor.

In the northern part of the shrublaml cadgibuts are replaced by Acacia
^(dicina and Acacia biveno^^a. Small drainage channels in this country are
bare of \‘egetation and meander through the samphire for a mile or so.

Thi'oughout this shrubland stock water can be obtained at no great depth.
Wooroo \V('1I on the Stock Route* and about three miles iidand from Wooroo
('r(*ek is an (*xample. When visited in duly the water level in this well was
about twelve* feet from the surface. This Avell is on the site of an old
aboriginal soak. Other w(*lis in the zone vary in de])th but no figures are
available*. On the coastal ]>lain, gre)und water is commonly too salt for stock
while in the spinifex country the* Avater though deeper down is, in most cases,
classed as “good stock water.’’

I>. S })inij'e.v PimUni.

The name “spinifex” is used locally for all species of Triodia; “pindan”
is a native word Avhich is used for the desert country by the ])astoralists.
This zone* can be* subdivid(*d as folloAVs -

( a ) Ti'ansitiona! ])has('
(])) Triodia ])hase.

( c) VJccfrachnc ])ha^e.

(a) The transitional phase is particularly clear along the Anna Plains
fi'ontage. It commences Avith a line of MeJaienca trees on the edge of the
treeless grey loam plain. The ground flora, includes small annuals such as
Sporobolus aiistralasicus and Kragro^stis Dielsil Avith occasional bushes of
Trianthema tnrgidifolia. AVhere the loam changes to the red sand the liora
also changes and Eragro:^tis eriopoda, Crotalaria Cnnninghamii, and Po-
lavisia icosandra re])lace the above mentioned species. This phase gives Avay
to a dense zone of Melaleuca lasiandra and Acacia transluceyis, Avhieh are
both .^'hrubs about three to five feet high, Avith occasional clumps of Triodia
pnujjens and Eragrostis eriopoda. After this the cadgibuts disappear and
the area can no longer be classed as transitional.

There aia* certain variations of the transitional phase. In some places
there is a band of Acacia salidna in others the Acacia trauslucens band is
missing. The change-over is narroAv and the transitional phase may only
be ten or 15 yards in depth. In other places it extends up to 50 but
this is not often.

(b) and (c) The spinifex ])indan has tA\'o definite phases. In the first
the pindan form of Triodia p)(ngeni^ is associated Avith a stunted form of
Bauhinia CunnhigJanuii. In the second Triodia is rej)laced }>y a similar
grass Plectrachne Schiri:ii and Bauhinia is replaced by small trees such as
DoUchandrone heieroph ffUa and Eacadgptus zggophglla.

The first of those phases is the nearer to the coast. Its inland boundary
is not very definite and the phase varies in depth. The Plectrachne coun-
tiy apparently runs out into the desert proper. It extended inland as far
as flu* authoi* trav(*ll(*d, i.t*., to a distance* of fifteen miles from the coast
at WTdlal. According to re]>ort it extends along tlu^ Rabbit Proof Fence at
h^t as far as the 121 degree* meridian.

Ecological Notes on the Vegetation of 80-Mile Beach. 103

n. Spinifex piiidan sliowiiig the assoeiatiou of slmibs and
tussock grasses. Acacia liolosericea, A. tiunida <ind FU'ctmclmc

Scliinsii.

Both phases of the ])indau carry various species of Acacia. These, ex-
ce]>t in the case ot Acacia i ranslnccn^^ which grows about two to three feet
high, an^ all small ti'(‘es. They commonly rmiu thickets in which only one
species is rejn'esented. The commoner species ai'(^ .1. hunida, A. l/(dosci'icca ,
-1. trachijcaypa, A. paclitjcarjai, A. corntcea, .1. >uipycss(j ami .!. stipnlifjeva.

In marked contrast to the coastal }>laiu arc the number ot shrubs which
are as^-ociated with the ])erennial tussock grasses. Thev include tlu' follow-
ing’: Ahattlo)/, l\alcoUti, .1. nidir/iin, Sidtt Icpida. ci. sj)inosa. Cahffhri.r m-
tcrstans^ Jlalgania Ulforalis, Piatclca annnoch.aris, .Id/v'ui/e tomr}it(}sa , Ct/ario-
■^tcgia Bunvgana , and Npw'.((Stlia- cJadoiricha. Larger forms are Luns/u gla-
tinosa, C. Sturtn var., Clerodendron tomentosum, Gardenia Paaioni and Da-
hoif^ia ITopwoodii and Htglohasimn fipathnlatum. Members of the Ib'oteaccae
are not numerically important. They include (h-erlUca pgramidalis, G. re-
fract((, (r, agrifoUa, IlaJica ma^croearpa and /Vraormm f(dcat<(. There art‘
many slender shrubs belonging to tlu‘ Leguminoseae. Kucalypts ar<‘ not
well represented. Tn most ])laces they are cum[)letely lacking fiami the
Triodia jihase while in tin* Plecti'achne country they gradually a})pear mor(‘
often as one goes farther inland. They either form small mallei' (lumps or
(kv('lop into small tri'cs from ten to tifteen feet high. The s] ;*<d<'s are
K'UraJg ptuH zggophglla and P. dichrowopidoia.

Plants of Pragrostis eriopoda ai'e often seen ainong-i the T) India tus-
socks. This grass is the most important s])ccies after Triodia and PJec-
Iracline since if replaces both when they are d(*stroyed and prcvtuli'd from
recovering through seedling development. Xormahy Eragrasii:< eriopoda is
]>resent in the association as a minor constitiu'iit.

Other grasses present are Amphij)ogon strictu^, (lirgt^opogon paUida;<^
Cyrnhopogon bombt/cina^^, Digitaria Brownei, Digitaria ctenantha. Panicinn
ci/mhiformCj I chnanthns aiistraliensis, Setaria s}irgensj . I ristida arenari<(,

104

Nancy T. Burbidge.

Knneapogon palUdiis, Eriaclnie pulchella aiul Sorg}iuyn phnnosnm. None of
these grasses is an important constituent.

The further one ]ienetrates into the desert the more numeioiis become
the long sand ridges. These, presumably, are similar to those described by
AVarburtoji (1875) and Carnegie (18f)S) in the country to the east. Gene-
ral observations suggested that the Avavla thickets grew between the ridges
or in open country while small plants of Banhinia Cunningliamii and small
trees such as Oicenia reticulata and Hakea ntacrocarpa grew on the ridges.

SrMMAKV,

An attempt has been made to describe the vegetation of the coastal
strip between Cape Keraudreri and Cape ^lississey. It has been shown that
there are two well defined ecological zones which are associated with cer-
tain soil differences.

On the coastal plain the. bioti(‘ factor has been responsible for the intro-
duction of Cenchrus ciliari.^ (Buifel Grass) especially on loamy soils. In the
desert country no introduced plant has as yet succeeded in establishing it-
self but where the normal tussock grasses are lost they are replaced by the
native species Kragrosti.9 rriopoda.

ACKNOWLEDGMENTS.

The author wishes to express her tlianks for assistance given by
the following; 'Mv. S. Mullins of Anna Plains Station, Mr. Spry of Nalgi
Station, Mr. Laev of Wallal Downs Station and Mr. Welch of Pardoo
Station. Without the co-operation of these the work could not have beo!)
carried out.

REFERENCES.

Carnegie, D. W.: Spinifex and Sand 1898.

Warburton, P. E.: Journey Across the Western Interior of .Australia :

Bbyozoa from the Wandagee and Xooncanbah Semes 105
(Pebmian) of Westebn Australia.

8.—BRY0Z0A FROM THE WANDAGEE AND NOON-
CANBAir SERIES (PERMIAN) OF WESTERN
AUSTRALIA.

Part One.

By Joan Ceockford^ iVLSc.

Read 14th April, 1942: Publish('d 24th March, 1944.

Communicated by Professor K. de C. Clarke.
('.ONTEX^rs.

Summary

Introduction

Description of Species : —

Family Fenestrellinidae Bassler.
Genus Fenestrellina d’Orbigny.
Fenestrellina horologia (Bretnall)

Fenestrellina disjecta sp. nov. ...

Fenestrellina ruidacarinata sp. nov.

Fenestrellina valentis sp. nov. ...

Fenestrellina columnaris sp. nov.

Fenestrellina lennardi sp. nov. ...

Fenestrellina cacuminatis sp. nov.

Genus Minilya gen. nov.
Minilya duplaris sp. nov.

Minilya princeps sp. nov.

Genus Polypora McCoy.
Polypora fovea sp. nov. ...

Polypora reiificis sp. nov.

Polyjiora woodsi (Etheridge)

Polypora muUiporifera sp. nov.

Polypora sp. nov. indet.

Acknowledgments

Bibliography

l^ige.

1()5

1()7

U)8

169

169

170

171
171

173

174

177)

176

177

177

178

I7i)

179

SUMMARY.

Fourteen species of Fenestrellinidae (Fenestrellina and Polypora), of
which twelve are described as new, and one neAV genus allied to Fenestrellina,
are described from the Wandagee Series of the North-West Basin and the
N'ooncanbah Series of the Kimberley District (Permian) of Western Aus-
tralia.

INTRODUCTION.

Bryozoa from the Permian and Carboniferous of Western Australia
wore first recorded by Hudleston in 1883, when he described Evactinopora
dendroidea Hudleston and E, crucialis Hudleston, and recorded Fenestellae,
Fenestella plehia McCoy, Protoretipora (sic) (FenesteUa) ampla Lonsdale
var., and Stenopora tasmaniensis Lonsdale from the ^'Carboniferous’^ North
of the Gascoyne River. Fenestella ("probably F. ampPP^) was recorded by
Foord (1890) from the Irwin River; Ilinde, in 1890, recorded Ilexayonella
dendroidea (Hudleston), and described Polypora australis Hinde and Bhoni-
hopora tenuis Hinde from the "Carboniferous” of the Gascoyne River Dis-
trict.

k 43/4 4-

Joan M. Crockfokd.

16 ()

H. Etlu'ridgo jnr. recorded Stenopora an<l Evacthiopora from Mt. Mar-
niion (1889); Hexayonella dendroidea and U. ("?) crncialis from Fossil Hill,
Wyndham River (1903); FejiestelJa fosstdtt Lonsdale from the Irwin River
(1907); and Evacthiopora crncialis and tAvo types of Steuopora (Stcnopora
spp. A and C) from Mt. Marmion, and Stcnopora sp. B from near Barra-
biddie (1915). Chai)man (1904) recorded Stcnopora leichardtii Nicholson
and Etheridge jnr. from the Irwin River District.

Bretnall (192(>) described L/fropora (?) crkosoidcs Pltheridge ms.^
Fenestella horologia Bretnall, E. affhtensa Bretnall, S nicorctepora ( ?)
meridianus Etheridge ms., Actomocladia ambrosoides Bretnall, Strcblotrt/jia
■marmionensis Etheridge ms., S. etheridgei Bretnall, Jthombopora niamillata
Bretnall, It. multigranulata Bretnall, and Coscininm (?) anstralc Bretnall,
and recorded ^thombopora tennis Hinde, from the Penno-Carboniferous of
the Gascoyne and Kimberley Districts. Miss L. Hosking (1931) recorded
Aetomodadia anib‘*'osoides, Coschmim (?) sp., Fenestella afflncnsa, F. horo-
logia, Rhombopora mamillata, R. muUigrannlata , St reblotr/rpa marinionensis,
and Snlcoretepora meridianus from the Wooramel River District.

Etheridge (1907 a) described some Permo-Carbonifei'ous bryozoa from
the Port Keats Bore, tliirty miles north of Fossil Head, Victoria River
estuary. Northern Territory, and discussed their relationships to AVesterii
Australian forms.

Gibb Maitland (1909), Raggatt (1930), and C'oiidit, Raggatt, and Rudd
(1930), have published lists of fossils, including Inwozoa, from the Permo-
Carbt>niferous of the North-West District, and Blatchfoi-d (1927) and AVade
(1937) from the Permo-Carboniferous of the Kimberley District.

The stratigraphical position of the Wandagee Series has been discussed
by Condit (1935), Condit, Raggatt, and Rudd (1930), Raggatt (1930), and
Teichert (1939, 1940, 1941); and the position of the Nooncanbah Series by
Wade (1937) and Teichert (1940, 1941). ■

The In’vozoa descrilied here are from the W^aiidagee Series of tlu' North-
West Basin, and from the Nooncanbah Series of the Kimberley District.
Fourteen species are recorded, and of these seven are known only from the
Nooncanbah Series, and two only from the AVandagee Series. The distri-
bution of the species is given iii Table I, and a comjmrison of their measure-
ments in Table TI.

Discussion of the age and affinities of the bryozoan fauna of the AATii]-
dagee and Nooncanbah Series is left until a later papei’.

DESCRIPTION OF SPECIES.

[^liylmn BKYOZOA Klirenberg’.

(bass (lYMNOLAEMATA Allman.

Order CR VPTOSTOMATA A'ine.

Family FENESTRELLINIDAE Bassler.

Genus FENESTRELLINA* d ’Orbigny.

Fenestrellina d’Orbigny, 1849, Revue et Magasin de Zoologie, 2 e Ser.,.

Tome I, p. 501.

Homonym: — Fenestella. Lonsdale, 1839, preoccupied for a pelecyp(xL
Fenestella Bolten, 1798.

* An application for suspension of the Rules of Zoological Nomenclature for the
generic name FeneiifeUa Lonsdale. 1839, has been submitted to the International Commis-
sion on Zoological Nomenclature (G. E. Condra and M. K. Elias. Journal of Paleonfoloov,.
Vol. 15, No. 4, pp. 565-566).

I5ry()zoa from thk Wandaoek and XooNCANiiAH Series
(Permian) ov Western Australia.

TABU*:

I.--l)iSTKIBrnoX OF SFKCIFS DFSCKi I5FD.

Wandagey Series.

Nooiicanl-ah S^^rie.s.

Other Localities.

(’alceolispongia Stage.

Linoproductus

Stage.

1.

2.

3.

4.

5.

(). !

1

7. !

s.

1

!). '

1

10, 1 11.

12 . j i.'i.

FenestreUina horo^ofjia (Hretnall)

+

4 .

4-

-F

“F

Gascoyne River Distri
R. ; Kampong Apn
to Baslcf* Beds),
Springsure District.

:*t : Callytharra Stage. W'ooramel

a, Bitauni, Noil Boewan (Bitaoeni
Timor : \'aucouvor Island, and

Queensland. .

FenestreUina disjerta sp. nov.

+

+

-F

FenestreUina ruidacarinata s]). nov.

4-

+ B.

FenestreUina valentis sp. nov.

t

4*

-F

FenestreUina columnaris sp. nov.

i_

+

-f

FenestreUina lennardi sp. nov. ...

4-A.

Feneshrllina cacuminatis sp. nov.

Minih/a dnjduris sp. nov.

1

A.

Gascoyne River Di.strict : Wooramel River DLstrict :

(?) Middle Prodiift.is Limestone. Salt Range, and
v8pringsure Di.strict, Queensland.

Minilya pri/icrpM sp. nov.

Folypora fovea sp. nov. ...

+

Folypora retifiei.f sp. nov.

4-

Folypora v:oodsi (Etheridge)

+

Stony Ck, Branxton. Mulbring, Jkdford. Clladulla (Branx-
ton Stage, Cpper Marine Series), N.S.W. ; l>c!ow

Sonoma Kd. C'rossing. Coral Gk. Bovvx*n R„ Queens-
land ; Marlborough. 'L'asmania : Callytharra Stage,

Wooramel lU : liitaoeni Beds, 'fimor, and Springsure

District. Queenslanil.

Folypora multiporifera sp. nov. ...

4-

4-

I

Folypora sp. nov. indet. ...

-F

1. Lower saiKiy i)urt of the Calceolispongia Stage, South side of Minilya River, 1|

to 14 miles Kast-North-Knst from Curdamu<!a Weil, Wandagec Station.

2. Low er half of the Calceolispongia Stage, ^ mile West of Coolkilya Fool.

3. Locality W.

4. Synclinc on Minilya River, A mile \\ est of Coolkilya Pool.

5. Uppermost part of the Cak^eolispongia Stage, North side of Minilya Kiver, 1

mile West of Coolkilya Pool.

(k Lowest bed of Linoprodu(;tiis Stage, East-VS'est striking fault block, North-W est
of North end of Wandagec Hill, on Telephone Line, just inside Woollies Pad-
dock, Wandagee Station.

i .

8 .

9.

10 .

11.

12 .
13.

28 chains North-East of Flag Station 12. Nalbia Pa<hlock, Wandagee Station.
Highe-st Pscudoga.strioceras horixon. 300 yds. East of South-West gullv f»f
Wandagee Hill. Mimgadan Pa<ldock. Wandagee Station.

Scarp 2 miles East ot Ohristma.s Ck. Homestejid.

0 miles East-North- Ea.st of Trig. Station (J2, St. George’s Range. Kimi)erley
District.

miles North of .Mt. Anderson, Kimberley DLtrict.

Keevio’s Well, 8 miles North of Mt. Andor.son Homesteacl, Kimberley District.
Mt. Marmion, Kimberley i)istrict ; A : Highest Beds of Nooncanbah Series,
S. side of ^It. Marmion ; B: Mt. Marmion (Fi7a47, .Australian Museum Coll.)

Bryozoa from the Wandagee and Nooncanbah Series
(Permian) of Western Australia.

Table II.— MEASUKKMENTS OF SPECIES DESCRIBED.

Specioj^.

I.orality.

Branches.

Fenestnilos.

Width of
Dissepiment
(mm).

Zooecia.

Nodes.

No. in
10 mm.

Width

(mm).

No. in
10 mm.

Length

(mm).

No. of
Row.s.

No. in
10 min.

No. per
Fenes-
tnile.

FenestreXlina horologia (Brotnalh

H milvs North of -Mt. Ander-
son Homestead

20-22

0-29-0-.38

16-18

0-29-0-52

O-I -0-29

2

37

2

High, in a single row, 0*23-
0*31 mm. apart.

Fenefflrellina disjecta sp. nov. ...

miles North of Mt. Ander-
son

12

0-2:M)-4

10

0*7 -0-94

0*14-0*24

2

40

4-5

Very high, elongated, 0*98-
1-17 mm. apart.

Fenesirellina ruidacarinata sp. nov.

9 miles East-North- Eivst from
Tri^. Stn. 02, St. George’s
Ra.

16

o-;«-o-38

10-10*5

0-6 -0*75

0-21-0-27

2

38

3-4

Hounded, contiguou.s, 0*13-
0 • 22 mm. apart.

FetiestreUina valentis sp. nov. ...

6^ miles North of Mt. Ander-
son

10

o-6;M)-7r)

10

0*52-0 -68

0-4 -0*48

o

33

3-4

Small, rather rounded, 0-24—
0 -4 mm. aj>art.

Fenesirellina columnaris sp. nov.

\ mile West of Coolkilya
Pool. .Minilya River

R>

o:i:M)-:js

14-15

0-44-0-5

0*22-0-3

o

37

3

Hiiih, thick, hlunt, 0-33-0*4
mm. apart.

Fenesirellina lennardi sp. nov.

South side of .Mt. Marmion

22

o- 2 i- 0 :i 2

:n

0- 17-0-25

about 01

2

31

1

Small, not well shown.

Fenesirellina caeuminalis sp. nov.

S miles North of .Ml. .Vnde?‘-
son HoiU(‘st<ad

22

11

0-.">2 0-63

about 0-12

42

3

Small, sharp. 0-24 -0-29 mm.
apart .

Minib/a dnplari'i sp. nov.

(>! miles North i^f .Mr. .Aiuf r-
.s<m

16 1!)

o-:j:t~o-4i

14-17

0-4 -0-51

0- 110-3

•>

-

33

2

Small, in two rows, o 13-0- 17
mm. a|Mirt.

Minibja prinreps sp. nov.

S miles North of .Mt. .Vndn-
son II«)me.'"t('ad

17

0 - n-o - 18

KMO-5

0-57 0-62

0-32-0-36

2

28

3

Sir.all, in two rows, 0- 13 0-21
mm. apart.

Pobfpora forea sp. nov. ...

H miles North of Mt. .Vnd'^r-
son IlomesK'nd

S-12

0 - l.")-0-7

6-6 ■ o

1- 15-1 -36

0- 17-0-29

4-5

31

4-5

Sn ail, irregularly «leveIo|x*d.

Polypora retijiris sp. nov.

mile West of C’oolkilya
Pool, .Minilya River

1.7-17

. „

0- IS-0-.77
L-.

9

0-67-0-S

0-41-0-6

3

33

4

Large, irregularly j)laml.

Polypora iroodsi (Ethcrid>r<*)

S miles Nortli of .Mt. An<l<*r-
son Homestead

10 1 I

0 • 0 • 62

8

0-68-0-92

0-46-0-63

3

31

4

Abundant, regularly pla<ed,
rather large.

J*olypora inulliporifera t^\). nov. ...

A mile West of (’oolldlya
Pool. .Minilya River

0-97-1 -I

1 •5-2-5

2 - i -5-6

0-46-0-6

5-6

22

6-14

Small, rath<*r abundant.

Polypora sp. nov. iiul(‘t.

Searp 2 miles East of (Mirist-
nu\s Ck. Home.stead

:i-4

1-2 -loo

1-5-2

3-3 -6-9

1-2 -1-6

about 8

9

9-15

? Not develojK*d.

I

- ,;^i

Bryozoa from the Wandagee and Nooncanrah Series
(Permian) of Western Australia.

1(>7

Inf HudibnUform or llabelUdo Fenestrellinidae, with tiro rows of alter-
natiuf) zooecia on the branches^ ewcejd before bifurcation, when the ninid)er
of rows is increased; rows of zooecia separated bp a mcdiam carina, jdain
or more fre(pientlp noth a single row of nodes {aca)ifho]/orcs) ; dissepiments
not cellalif crons ; internal stractnre and rererse snrfarr as for the f<fmihp

Fenestrellina horologia ( llrclnall 1 (Plato I., figs. (>.)
Fenestella horologia Bi’olnall, 19-(5, dosc-riptioiis of somo Woshaai Austra-
lian Fossil Poiyzoa, W.A. (leol. Sarr. HnU. SS, p. lo, pi. 1, tig. ti.

[non] Fenestella horologia Bri'fnall, ilosking, 1991, Fossils froni tho \V(Kn‘a-
inel Distriot, W.A., Jonr. Hop. Sor., ll’..!., A”ol. XVII., p. 19, pi. 1\.,
fig. 9.

Fenestella parvinsenia Basslor, 1929, Ponnian l^>ryozoa of Tinioi', Pa! lontolo-
gle ro}i Timor, Liof. XVI., XXVllL, p. 7(i, pi. (’(WLl (IT), tigs. 8-19.

Fenestella parrinsenia Basslor, Martin, 19,92, Do Paloontolouio an vStrati-
graphio van XcMlorlandsoh Oost-lndio, Bryozoa, p. 991.

Fenestella parvinsenia Basslor, Fritz, 1992, Ponnian Bryt)Z(»a from Vaiu-om'or
Island, Trails. Hop. Soe., ('anada, Ser. 9, Vol. 2t), S('ct. I\’., p. 9t).

Fenestrellina parrinsenia (Basslor), Flias, 1997, Slratigraphii- Siguitioanoo
of Sonio Lato Paloozoio Foin'strato Bryozoans, Jonr. Paleontologp, Vol.
11. No. 4, p. 914.

Fonosti'(dlina irith (wo zoeeeia to a fenest rale ; earina Ion'; nodes sharp,
relativelp high, abont two to a fenest rale ; fenesl rales honr-glass shaped.

Horizon and looality: Xoonoanhah Sorios; Ko(*vi(‘'s \V('ll, (dght miles
north of Mt. Andorson Ilonu'stoad, Kimborloy Distriot (('oil. A. Wado).

The form of tlio oolony is not shown; thoro aro f)-om 1(> to IS I'onostrnlos
vortioally, and from 20 to 22 branohos horizontally, in 10 mm. Tho branohos
aro straight, from 0-29 to 0*98 mm. in widtli, and show a slight, i-atlu'r
roundod oarina, with sharp, ivlativoly high nod(‘s in a singlo row, at intor-
vals of from 0 29 to 0-91 mm.; tho bases of those nodes are slightly elon-
gated i)arallol to tho length of tho bram-hos. In tho older parts of tlu* colony
supporting rootlets may bo developed from tho oollulitorous suriaoo, and
tho thioknosH of the bi-anolios (normally about 0-9 mm.) may bo gi-oatly
inoroasod by extra deposits of oaloium oarbonato: the nodes boeonu' very
muoh thickened, and may appear bifid or tritid near tho top, and tho aper-
tures b(M'onio closed. The sides of tho ])ranohos, on which tho apertures aro
placed, aro slightly (lattenod, but tho apertures project into tho fonostrnlos;
they aro circular, from 0-1 to 0-19 mm. in diameter, amt where they are
w('ll preserved show about eight fine radially directed ribs; tlu'se, however,
were v(*ry easily iemo\'ed bv weathering; the peristomes ar<‘ distinct, and
ai ‘0 best developed on th(‘ side towards the Uaiestrules. 1 w<t apertures o<'cur
in the length of one fem'striile and one <liss(‘piment ; these art* nsnatly V(*ry
regularly arranged, one being phn't'd o])positc the end ot t'ach disst'pimenl .
and one at the centre of each fenestrule -the projection of these at>ertim's
into th(* fenestruh's gi\es tlu-m tlndr characterislit-, hour-glass shape;
a less regular arrangement may be develoi>ed lor a it'w lenestrnles.
The distance between the centres of successive aiu'rtures is from
9-21 to 0 9.4 mm. (avt'rage 9-27 mm.}, but is generally betwe(‘ii 9-27) and

* TliivS species is beiiij; revised, in r st'i.nraio paper, from the type mtitorial in the
collection of the Western Anstruliun Geological Survey, with which the specimens here
described have been compared.

1(J8

JOAX ^I. CROCKFOm).

0-3 mm.; about thiry-seven apertures occur in 10 imn. The fenestrules
are from 0-29 to 0-52 nmi. in length and from 0-17 to 0-4 mm. in width;
the width of the dissejiiments is from 0-1 to 0-29 mm., but is generally
less than 0-2 mm., and the length of one fenestrule and one dissepiment
is from 0-49 to 0-65 mm. (average 0-57 mm.). The dissepiments, when'
they are well-preserved, may show two or more fine transverse ridges, and
the whole of the celluliferous surface is covered by very tine tubercles. On
the reverse surface both branches and dissepiments are evenly rounded, and
the dissepiments may be as thick as the branches; the outermost layer is
finely granular, and the backs of the branches may be ornamented by fine
nodes, thoug’h these are not always developed over the whole of the reverse
surface, and are easily worn away. The inner layers of the reverse surface
show fine longitudinal striae. Bifurcation occurs usually at distant intervals,
though it may be frecjuent; increase to the three rows of zooecia occiu’s im-
mediatelv before branehing:.

Remarks: Fenesfr€lli}ia horologia was described by Bretnall from the
Gascoyne River District; it occurs as low in the Permo-Carboniferous as
the Callytharra Stage; and occurs also in Queensland (Consuelo Ck., two miles
above Cattle Creek, S]'»ringsure District; Reid, 1930, p. 157, locality 9).
It appears to be the same as Fenesirellina parvinscula (Bassler), which occurs
in Timor in both the Bitaoeni and Basleo beds. F. parvinscula has also been
recorded from the Permian of Vancouver Island.

Fenestrellina disjecta sp. nov.

Plate 1, Fig. 2; Text-figure 1 F.

Holotvpe: Specimen 2793a, University of Western Australia Collection.
(Coll. IT. W. B. Talbot.)

Horizon and locality: Xooncanbah Series; fiV'g Xorth of ]Mt.
Anderson.

Fenestrellina with four to five zooecia to a fenestrule ; slight carina;
verg high nodeSj one occurring in the length of each fenestrule. and usually
placed opposite the junction of a branch and. a dissepiment.

The form of the colony is not shown ; there are 12 branches horizontally,
and 10 fenestrules vertically, in 10 mm. The branches are straight, from
0-33 to 0-4 mm. in width, and shoAV a slight median carina, bounded by a
depression on each side. Very high nodes, Avith their bases elongated parallel
to the length of the branches, occur at inteiwals of from 0-98 to 1-17 mm.,
and are generally jdaced opposite the junction of a dissepiment Avith a
branch; in well preserved specimens the height of these nodes is up to
0-25 mm., but they usually appear very much smaller. The apertures are
circular, 0 08 mm. in diameter, and are surrounded by distinct peristomes.
Four, less often fiA'e, apertures occur in the length of one fenestrule and one
dissepiment; the distance betAveen the centres of successive apertures is from
0-21 to 0-28 mm. (average 0*25 mm.') and about forty apertoes occur in
10 mm. The fenestrules are oA'al, from 0-7 to 0-94 mm. in length, and from
0-29 to 0-54 mm. in Avidth; the dissepiments arc from 0-14 to 0-24 mm. in
AAudth, and the length of one fenestride and one dissepiment is from 0-89
to 1-lfi mm. On the celhdiferous suiTace the dissepiments are slightly
rounded and shoAv four or five fine ti*ansA*erse ridges and grooA'es. On the
reverse surface both branches and dissepiments shoAV numerous fine longi-
tudinal striations AAliere they are slightly weathered. The dissepiments are
distinctly thinner than the branches. At their junction Avith the dissepiments

Hhvozoa krom the VV'am)A(;kk and Xooncanraii Skrirs
( Pkrmian) of Western Austraei a.

the branches show fairly large, rather indistinct tubercles on the reverse
surface. The lateral margins of each branch are angular. Bifurcation may
occur at intervals of about T mm., but is usually much less freqiumt; increase
to thi’('e rows ofc‘ zooecia occui's immediately before branching.

Bemarks: This s]>ecies is distinguished from described species of similar
size by its large, widely spaced nodes.

Fenestrellina ruidacarinata sp. nov.

Plate 2. Figs. 1, 2.

llolotype: Sj)ecinien 2707a, ruiv(u*sitv of Western Australia Collection.
(Coll. IT. W. B. Talbot).

.Horizon and locality: Nooncaubah Series; !) ni. Fj-N-E. from Trig. Stn.
Cr2, St. George’s Itange.

Fenestrellina with three to four zooecia to a fenestrule; carina low^
nodes rounded^ contkjxious, five to six in the length of each fenestrule.

The form of the colony is not shown; there ai‘e K> branches horizontally,
and 10 to lO o fenestrules vtu'tically, in 10 miu. Tlu‘ branches are sti*aight,
and are rather rounded on the c('lluliferous surl‘a('e; the apertures are ])laced
on the sloping sides of the bi’anches, but do not i)roject into the fenestrules;
the width of the branches is from 0-23 to 0.38 mm. The carina is almost
obsolete, and is com])letely covered I)y the nodes, Avhicli are low, rounded,
and contiguous, and which vary in diameter, though they do not vary in
height, except as a result of weathering, which may remove all traces of
the divisions between adjacent nodes and leave only a slight ridge along the
centr(‘ of the ]>ranch. Five to six nodes occur in the length of each f(mes~
trule; the distance between the centres of successive nodes is from 0 1 3 to
0-22 mm. The apertures are circular, 0-11 mm. in diameter, and are sur-
rounded l)y distinct, but only slightly raised, ])eristomes; three apertures
occur in the length of each fenestrule, and an (>xtra one may occur opimsite
the dissepiment; the distance between the centres of successive a)KU’tures is
from 0-24 to 0-3 mm., and about thirty-(‘ight Jipertures occur in 10 mm.
The fenestrules are oval, j’rom 0-(i to Own mm. in length, and from 0-24
to 0-35 mm. in width, and tin* dissepiments av(' from 0-21 to 0-27 mm. in
width. On the celluliferous surface each disse])iment shows one or two
strong transverse furrows. The reverse surface is not shown. Bifurcation
occurs at intervals of more than 5 mm.; increase to three rows of zooecia
occurs within one fenestrule before branching.

Kemarks: The numerous low, contiguous nodes distinguish this form
from any describ(*d Permian specie's. Fenesirellina inaequalis (Ulrich), from
the U])per Coal Measures of Illinois, is similar in a])pearanc(* ; it is, how-
ever, a large'r s])ecies, wdth rather more zooecia to a fenestrule, and shows
smaller and much more* numerous nodes.

Fenestrellina valentis sj). nov.
Plat(* 1. Fig. 4.

Holoty[)e: Specinu'u 2703c, Uui\(*rsity ol‘
(Coll. H. *W. B. Talbot).

Horizon and localitv: Xhauicanbah Sei’ies

Western Australia Collection.
()V 2 m. North of !Mt. Ander-

son,

170

JOAX M. Crockford.

Feiiestrelliuil with three to four zooecia to a feuestrulej very broad
bra'nches ; rounded carina with small nodes.

There are 10 branches horizontally, and 10 fenestrules vertically, in
10 imii. The branches are very broad, from 0-03 to 0-75 mm. in width, and
show a median, rounded, relatively low carina, with rather small, rounded
nodes, placed from 0-24 to 0-4 mm. apart. The apertures are circular,
0 13 mm. in diameter, and are placed on the tlattened sides of the branches;
no peristomes are shown. From three to four apertures occur in the length
of one fenestrule and one dissepiment; th(‘ distance between the centres of
successive ai)ertures is from 0-29 to 0-35 mm., and about 33 apertures
occur in 10 mm. The fenestrules are oval, from 0-52 to 0-68 mm. in
length, and from 0-3(i to 0-44 mm. in width; the width of the dissepiments
is from 0-4 to 0-48 mm. A cast of the reverse surface shows that both
branches and dissepiments are evenly rounded, and that they are of the
same thickness (about 0-4 mm.). On tlu‘ c(41uliferous surface the dissepi-
ments show numerous tine transverse striae Avhen they are very slightly
weathered.

Kemarks: The broad, tlattened branches and dissepiments distinguish
this form from associated species.

Fenestrellina columnaris sp. nov.

Plate 2, Fig. 3; Text-figure 1 F, G.

Holotype; S[>ecimen 20949, I’niversity of Western Australia Collection.
(Coll. C. Teichert and H. Coley.)

Horizon and locality: Lower half of the Calceolis))ongia Stage of the
Wandagee Series; Syncline on Minilya H., mile West of Coolkilya Pool.

Fenestrellina with three zooeeda to a fenestrule; carina slight, nodes large
ayid very high.

The colony Avas probably infundibuliform ; the holotype is a large, very
much folded expansion. There are 10 l)ranches horizontally, and from 14
to 15 fenestrules vertically, in 10 mm. The branches are straight, from
0-33 to 0-38 mm. i!i Avidth, and shoAv a slight median carina, Avhich bears
a single roAv of high nodes, placed from 0-33 to 0-4 mm. ainirt. These nodes
are slightly elojigated along th(‘ carina at their bases, but above this they
are rounded in cross section, and terminate bluntly; their height is up to
0-17 mm., and their diameter at the top may be as much as 0-2 mm., though
it is generally less. The apertures are circular, 0-13 mm. in diameter, and
are placed on the sloping sides of the branches, but do not project into the
fenestrules; }>eristomes are not developed. Three apertures occur in the
length of each fenestrule, and one of these may be placed opposite the end
of a dis.sepiment ; the distance between the centres of successive apertures
is from 0-25 to 0-29 mm., and about thirty-seven apertures occur in 10 mm.
Bifurcation of the l)ranches may occur Avithin 5 mm. ; increase to three i‘ows
of zooecia occurs immediately before branching. The fenestrules are oval,
from 0-44 to 0-5 mm. in length, and about 0-25 mm. in Avidth ; the Avidth
of the dissepinu'iits is from 0-22 to 0-3 mm. On the reverse surface both
branches and dissepiments are evenly rounded, although after .slight Aveather-
ing they appear angular; the dissepiments are as thick as the branches —
about 0-75 mm.; the outermost layer of the revers(‘ surface is finely granular.
Rootlets, which are often forked, and are up to about 8 mm. in length, may
])e developed from the reverse surface.

Bryozoa from the Wand AGEE and Nooncanbah Series 171
(Permian) of Western Australia.

Remarks: From Fenestrellina horologia (Bretnall) this species is
distinguished by its larger size, by having more zooecia to a fenestrules,
and by its large nodes. Fenestrellina jjulchrador salts (Bassler) is a larger
s])eeies, and does not show large nodes.

Fenestrellina lennardi sp. nov. (Text figure lA.)

Holotvpo: Specimen 20f)48a, University of Western Australia Uollection.
(Coll C. Teichert.)

Horizon and locality: Highest beds of the Nooncanbah Series; South
side of Mt. Marmion, Kimberley District.

Fenestrellina with one aperture to a fenestrule; carina slight; nodes
small.

The form of the colony is not shown; there, are about 31 fenestrules
vertically, and about 22 branches horizontally, in 10 mm. The branches are
straight and flattened, and show a slight, rounded, rather wavy carina; small
nodes are developed, but are not well shown. The branches are relatively
very broad, being from 0-24 to 0-32 nmi. in width; the zooecial apertures
are circular, 0-1 mm. in diameter, and are suiTounded l>y slight jieristomes;
they are usually very regularly placed, one occuring opposite the end of
each dissepiment. The distance between the centres of successive apertures
is from 0-29 to 0-38 mm., and about 31 occur in 10 mm. The fenestrules
are oval, from 0 17 to 0-25 mm. in length, and generally about 0.21 mm.
in width; the width of the dissepiments is about 01 mm. The
reverse surface is not shown. Bifurcation occurs at relatively distant inter-
vals, and increase to three rovs of zooecia occurs just before branching.

Remarks: The small size of this species, and the arrangemenf of the
zooecial apertures, separate it from associated s])ecies.

Fenestrellina cacuminatis s]>. nov. (Text figure IB.)

Holotype: Specimen 20944a, I'niversity of AVestern Australia Collection.
(Coll. A. Wade.)

Horizon and localitj' : Nooncanbah Series : Keevie’s Well, 8 miles North
of Mt. Anderson Homestead.

Fenestrellina with three zooecia to a fenestrule; carina slight; nodes
small, sharp, evenly spaced.

The form of the colony is not shown ; there are 14 fenestrules vertically,
and 22 branches horizontally, in 10 mm. The branches are straight, about
0-25 mm. in width, and the carina is formed principally by the junction of
the sloping sides of the branches; the nodes are small and sharp, but not
high, and are placed from 0-24 to 0*29 mm. apart. ,'The apertures are
circular, 0 08 mm. in diametei', and are ])Iaced on the sloj)ing sides of the
branches; slight peristomes are developed, and where they an' well-i)reserved
the apertures project slightly into the fenestrules. The distance Ix'tween the
centres of successive apertures is from 0-22 to 0-27 mm., and about 42
occur in 10 mm.; there are three apertures to a fenestrule — one of these
may be placed opposite the end of a dissepiment. The fenestrules are oval,
from 0-52 to 0-03 mm. in length, and from 0-17 to 0-25 mm. in width; the
dissepiments are about 0-12 mm. wide. On the obv^erse surface the dissepi-
ments when slightly worn, show a few transverse grooves. The reverse
surface is not shown.

Remarks: This species is distinguished from Fenestrellina pulchradorsalis
(Bassler) by its narrower branches and dissepiments, and smaller size. F.
affluensa (Bretnall) is a larger species.

172

JOAX Crockford.

Genus MINILYA gen. nov.

Fenestrellinae in which the branches show two rows of alternating
zooeckiy one on each side of a slight median carina; nodes small, in two
rows on the carina^ placed so that one node is lateral to each zooecial aperture;
zooecia sub-triangular ; structure of the reverse surface as in Fenestrellina.

Range : Upper Pennsylvanian to Permian.

Genotype: Minihja duplaris sp. nov.

A. Fenestrellina lennardi sp. nov.: Celluliferous surface of the holotype
(No. 20948a, Univ. of Western Australia), x 10.

B. Fenestrellina cacuminatis sp. nov. : Celluliferous surface of the holotype
(No. 20944a, Univ. of Western Australia), x 10.

C. Minilya duplaris sp. nov.: Portion of a thin section of a specimen from
the Calceolispongia Stage of tlie Wandagee Series (Specimen in the University
of Western Australia Collection), showing the double row of nodes and the
arrangement of the zooecial apertures; the section has cut just above the level
of the dissepiments, x 10.

D. Minilya duplaris sp. nov. : A thin section of the same specimen, showing
the shape of the zooecia, and, on the left hand side, some of the tubercles charac-
teristic of the reverse surface of this species, x 10.

E. Fenestrellina disjecta sp. nov. : Lateral view* of part of one branch of a
topotype (Specimen 2793b, University of Western Australia Collection), x 10.

F. G. Fenestrellina columnaris sp. nov.: Lateral views of two branches of
the holotype, x 10.

{Camera lucida diagrams.)

Species belonging to this late Palaeozoic group have been de-
scribed from the Upper Pennsylvanian of Texas and Kansas, and
from the Permian of India, Timor, Kansas, Texas, Nebraska, and Eastern
and Western Australia. A list of the measurements of some of the species

BkYOZOA from THK WaNDAGKE and N()(JN'CANBAH

(Pkrmian) of Westfhn Austuadia.

17 :^

belonging here has been given by Elias (1937, p. 324). The following are
amongst the species included in this group : —

Fenestella geminanoda Moore (Upper Uennsylvaiiian to LoAver
Permian).

Fenestella bispimdata Moore (Upper Pennsylvanian).

Fenestella hinodata Condra (Upper Pennsylvanian).

Fenestella conradi var, compactilis Condra (Upper Pennsylvanian
to Lower Permian).

Fenestella kukaensis Bassler (Permian).

Fenestrellina hitnherculata Crockford (Permian).

Fenestella perelegans Meek, Waagen and Pichl (not Fenestella
perelegans Meek), (Permian).

^Fenestella jabiensis Waagen and Pichl (Permian).

Minilya duplaris sp. nov. (Permian).

Minilya princeps sp. nov. (Permian).

Minilya duplaris sp. nov.

Plate I, hgs. 5, 7 ; Text-figure 1 C, D.

Fenestella^ horologia Bretnall, ITosking, 1931, Joar. Roy. Soc. W.A.^ Vol.
XVII, p. 13, pL IV, fig. 3. '

[non] Fenestella horologia Bretnall, 1926, IV.A. Geol. Surv. Bull, 88, p.
15, pi. I, fig. (1.

[cf.] Fenestella perelegans Meek, Waagen and Pichl, 1883, Pal. Indica, Ser.

XIII, Vol. I, p. 777, pi. LXXXVII, figs. 1-3.

[non] Fenestella perelegans Meek, 1871, in Ileyden’s Final lU'port on N('-
braska, p. 153, p\. VII, 6g. 3.

Holotype: Specimen 2793d, University of Westeni Australia Collection.
(Coll. H. W. B. Talbot).

Iloiizon and locality: Xooncanbali Seri(‘.<, miles North of Mt.

Anderson.

Minilya with two zooecia to a fenestrule; carina slight; nodes sharp,,
relatively high, in two roies on the C((rina; reverse surface tuberculate.

The form of the ('olony is not shown: there are from 16 to 19 branchi's
horizontally, and from 14 to 17 fenestrnles A'ertically, in 16 mm. The
branches are sti’aight, fi'om 0-33 to 0-41 mm. in width, and show a slight
median carina, which may be produced upwards in a very thin, zig-zag'
ridge betAveen the node>, Avhich are in tAA'o rows and ‘cWo. alternating; the
nodes, where they are \vell pres(‘rved, are sharp and relatively high, and
each node is placed lateral to an aperture; adjacent nodes are from 0-l!l to
0*17 mm. apart. The a])ertur(*s are circular, al)out 613 mm. in diameter;
very thin ])eristomes are developed, but these are usually broken away. Tin*
apertures are placed on the steeply sloping sid('s of the brances, and pro-
ject into the fenestrules; the distance betAveen the centres of successive
apertures is from 0-27 to 6-35 mm. (average 0-3 mm.), and about 33
occur in 10 mm. There are two a])ertures in the length of one fene-

* Til 1915 Frederiks (pp. 47-48) described a new sub-genus of Fenestella, Fenestepora,
with three species, Fenestepora jabiensis (Waagen and Pichl), Fenestepora foraminosa
(Eichwald), and Fenestepora retiformis (Schlotheim) . The species considered by Frederiks
to be identical with Fenestella jabiensis Waagen and Pichl is a different species : Fenes-
tepora is considered to be a distinct genus, and Fenestepora jaln^nsis Frederiks [not Waagen
and Pichl] has been chosen as genotype by Bassler (1935).

Fenestepora Frederiks is distinguished from Fene.streUina d’Orbigny by the presence-
of a row' of small cellules (as distinct from nodes) on the carina of each branch.

174

Joan M. CiiOCKFOKo.

show five or six
so thiek as the
emisepta are de~
1 I); tliev show

strule and one diss(*pinu>nt, and these are j»-enerally regularly plaeed, with
one op])osite the end of each (!isse])iinent, and one at the centre of each
fenestrule, hut one S])eciinen shows slight irregularities in the placing of the
apertures. The feneslndes are hourglass-shai>ed or oval, and are from
0-4 to 0-hl mm. in length and from 0 14 to 0-25 mm. in width ; the dissepi-
ments are from 0-11 to 0-J mn'i. in width, and the length of one fenestrule
and one dissepiment is from 0-51 to O-Td mni. The reverse surface is not
shown in specimens from tin* type locality, hut si)ecimens from the highest
beds ol the Nooncanhah seu'ies at Ml. IMarmion, and from tlie Calceolis])ongia
Stage of the Wandagee Sen-ies on tlu‘ Minilya Kivei- show that the reverse
surface of the hranclies is ornamenttHl hy \ery numerous, large, irregularly
t>laced tuhercles; the inner layers of the r(‘vei*se surface
line longitudinal striae'. Tlu' dissepiments are not (piiti
hranclies. Sections show that hoth superior and inferioi'
veloped; the shape of tlu' zooecia is shown in Text-tigun
a greater resemblance to those described by Nikiforova (lOJJ, ]>. 13) for
the FenestreUi^m dov.aiea grout) thosi' of other forms. The tubercles

of the reverse surface show in sections as rounded areas of lighter colour.
Bifuri'ation generally occurs at infreijuent intervals, but may occur within
3 mm.; increase' to Ihrei' rows ol zoot'cia occurs imnu'diately before branch-
ing.

K(*marks: 1 his species is probably the form described by Miss Ilosking
(1031) fj'om tin' M ooramel Rivei' Disti'ict as Fenestella horologia Bretnall,
but it differs from F. horologia in Inning a double, instead of a single, row
of nodes. It occurs in the Sjiringsure District, Queensland (Consuelo Ck.,
two miles abo)e Cattle Ck.: Heid, 1930, p. 157, locality 9). Fenestella
<‘onmdi var. compactdis (’ondra, fi’om the Upper Pennsylvanian and
Lower Pei’inian of Kansas and Nebraska, is dift'erentiated by having
less regularly arranged ai>ertures and smaller nodes, which may
not always be placed in {\yo r<iws, as they are shown in Condra^s figures;
the measurements and figure's given by Condra for thi.s species do not al-
ways corres])ond. Fenestella bispinnlata Moore, from the Upper Penn-
sylvanian of Kansas, is a smaller species, and diffei's in the a])])earance of
the reverse surface. The s])ecif-s recoreh'd as Fenestella pereJegans Meek
from the Middle Pi'oductus Liim'slone of the Salt Ranges by Waagen anti
Pichl is slightly finer than this species, as was noted by Miss Hosking, and
possibly shows a rather higher carina : tlu* reverse surfaces of the specimens
examined by AVaagen and Pichl wt'i’c very much weathered; it is possible
that these two species art* identical. Fenestrellina pereJegans (Meek), from
the (’oal Mt'asurc's ot Nebraska, ]>ossessed only a single row of nodes on the
t'arina.

Minilya princeps sj). nov.

Idate 1. Pig. 1.

Holotyju*: S])ecim(*n 20945, I'niversity of Western Australia Collection.
(Coll. A. Wade.)

Horizon and locality: Nooucanbali Series; Keevie’s AVell, 8 miles north
of Mt. Anderson Homestead.

Minilya idth three zooeeda to a fenestrnle ; carina almost obsolete; nodes
small, in tico rows on the carina.

The form of the colony is not shoAvn ; there are from 10 to 10-5
fenestrules vertically, and 17 branches horizontally, in 10 mm. The branches

175

Bryozoa from the Wandagee and Nooncanbah Series
(Permian) of Western Australia.

aiT straight, and are rounded on
0-41 to 0-48 iniu. in width. The

the eelluliferous surface; they are from
Carina is broadly rounded and is ])oorly

defined, and shows two rows of rather small nodes, which regularly alternate;

each nod(‘ is placed lateral to a zooecial aperture; the distance between
adjacent nodes is from 0 13 to 0-21 uim. The apertures are circular, 0-13
mm. in diameter, and are ]>laced on the sloping sides of the branches: they
may ])roj(*ct very slightly into the fenestrules ; the i^eristomes are very ])oorly
defined. There are usually two ai>ertures in the length of each fenestrule, with
an extra oiu* opposite each dissepiment; the distance between the centres of
successiv(‘ a])('i'tu]‘es is al)out 0-35 mm., and twenty-eight apertures occur in
10 mm. The fenestrules are oval, fi‘om 0-57 to 0-02 mm. in length, and
about 0-2 mm. in width; the . Avidth of the dissepiments is from 0-32 to
0-3G mm. On tlu^ eelluliferous surface the dissepiments are rounded, and
are ('overed by a thick, finely granular deposit; they are not dejn’essed very
much below the surface of the branches. On the reverse surface the branches

and disse])iments are of the same thickness; both are evenly rounded; the
outermost layer is finely granular. Bifurcation occurs at infre(|uent inteiA'als,
and increase to three roAvs of zooeeia occurs immediately before branching.

Remarks: This species differs from Fenestella jahipusi^ Waagen and
l^ichl, from the Middle and Opper Productus Limestone of the 8’alt Ranges,
in having fewer zooeeia to a fenestrule, and a less well defined keel, and in
having the dissepinumts on the eelluliferous surface nearly level with the
branches; and from Fenestella hinodata Condra in having feAA'er fenestrules
in 10 mm., and much more widely spaced zooeeia. Minilya duvlaris s}). nov.
is a finer form, and shows fewer zooeeia to a fenestrule.

Geinis POLYPOEA McCoy.

Polypora McCoy, 1845, Synopsis of the Carboniferous Limestone
Fossils of Ireland, p. 207.

Genoty]K‘ : Polypora dendroides McCoy, 1845.

Jnfnndibuliform or flahellate Fenestrellinidae , ivith the zooeeia arranged
in three or more rows on the hranclies, except just after bifurcation, ivhen
ftnly two rows may he ]>resent ; nodes (acanthopores) usually present, hut
carina not usually developed ; dissepiments not eelluliferous ; internal structure
and reverse surface as in Fenestrellina.

Polypora fovea sp. nov.

IMate 3. Figs. 1, 3.

Holotvpe; Sp(‘cimen 20047, Fniversitv of Western Australia Collection
tColk A. Wade.)

llorizoii and locality: Xooncanbah Series; Keevie’s Well, 8 miles north
of lUt. Anderson Homestead.

Polypora with four to five rows of zooeeia, and four to fire zooeeia to a
fenestrule : nod.es small, infrequently developed ; branches and dissepiments
narrow.

There are 8 to 12 branches horizontally, and 0 to 6.5 fenestrules verti-
cally, in 10 mm. The branches are straight and are relatively very narrow,
being from about 0-45 mm. in Avidth where there are four, to as much as
0-7 mm. Avhere there are five, roAvs of zooeeia; they usually show four to
five rows of zooeeia, with three — less often tAvo — for one or tAvo fenestrules
aft(*r, and five to six just before, branching. The apertures are sub-circular,

17()

JOAX M. Ckockfokd.

0-11 by 0 06 mm. in diameter, and are surrounded by well-developed
peristomes; they may be closed by a calcareous plate, frequently showing a
small central perforation; the whole of the celluliferous surface shows strong
ridges and grooves around and between the ai:>ertures. There are five, less
often four, apertures to a fenestrule, the distance between the centres of
successive apertures being from 0-25 to 0-35 mm.; about thirty-one apertures
occur in 10 mm. Small nodes are irregularly developed. The fenestrules are
sub-rectangular, from 1-15 to 1-36 mm. in length, and from 0-3 to 0-95 mm.
in width; the dissepiments, which expand only slightly at their junction
with the branches, are from 0-17 to 0-29 mm. in width. On the celluliferous
surface the dissepiments are evenly rounded, and each shows four or five
strong transverse ridges and grooves. On the reverse surface both brancht
and dissepiments are rather sharply rounded; the branches are rather thicker
than the dissepiments, and show numeroxis fine longitudinal striae w^hen they
are very slightly Avorn ; a roAV of fine tubercles may occur across the back of
a branch at its junction Avith a dissepiment, but these are not ahvays
develope'd. The dissei)iments shoAv fine transverse ridges and grooves. Bifur-
cation of the branches occurs at intetwals of 5 mm. or more.

Remarks: Pohjpova virga Laseroii, AA'hich occurs in the Upper Marine
Series in New South Wales, is a larger form, with usually three rows of
zooecia.

Polypora retificis sp. nov.

Plate 2. Fig. 4.

llolotype : Specimen 20951, L niA'ersity of estern Australia Collection.-
(Coll C. Teichert and H. Coley).

Horizon and locality: Calceolispongia Stage of the Wandagee Series;
Syncline on iNIinilya Ri\'er, half mile AA est of Coolkilya Pool.

Polypora with three roivs of zooecia, and four zooecia to a fenestmle;
nodes veri/ large, Imt infrequentlg developed.

Tlie form of the colony is not shoAvn; there are 0 fenestrules vertically,,
and 15 to 17 branches horizonttdly, in 10 mm. The branches are straight
or slightly [lexuous on the obverse surface, and normally sIioav three roAVs
of zooecial apertures, Avith four just before, and tAvo for about two fenes-
trules after, branching; there is a slight carina where only tAvo rows ot
apertures occur. Large nodes about 0-22 mm. in diameter are iriegularh
and infrequently developed, generally in the mid-line of the branch. There
are four apertures in the length of one fenestrule and one dissepiment,
either one or two of these being placed opposite the end of the disse})iment ;
the distmice between the centres of successive apertures is from 0-24 to 0-.-.S
min., and about thirty-three apertures occur in 10 mm. The apertures are
circular, about 0-11 mm. in diameter, and show thin peristomes; over a
large part of the surface the apertures have been enlarged, and the peri-
stomes remov(’d, by Aveathering. The apertures may encroach on the end:^
of the dissepiments, but extra cells do not normally occur; in one place
two branches each shoAving tAVo rows of zooecia coalesce tor a short distance
after their separate bifurcations. The fenestrules are oval on the cellu-
liferous surface, and are from 0-67 to 0-8 mm. long, and from 0-33 to 0 5
mm. Avide: the Avidth of the dissepiments is from 0-41 to 0-6 nnn., and the
length of one fenesti’ule and one dissepiment is from 1*16 to 1-38 mm. On
the reverse surface both branches and dissejiiments are rounded, and the
fenestrules appear round or OA'al; the dissepiments are generally slighth

Bryozoa from the Wandagee and Nooncanbah Series
(Permian) of Western Australia.

177

thinner than the branches j the outermost layer of the reverse surface is finely
granuiarj but when it is weathered fine concentric striae are shown. On the
celluliferous surface the branches are from 0*48 to 0-57 mm. in width where
there are three^ from 0-3 to 0-38 mm. where there are two, and about 0-85
mm. where there are. four, rows of zooecia. Bifurcation may occur within
5 mm.

Remarks: Pohjpora tcoodsi (Etheridge) is differentiated by its far more
numerous, and much smaller, nodes.

Polypora woodsi (Etheridge).

Plate 3, fig. 2.

Frotoretepora cimpla var. ivoodsi Etheridge, 1892, Geology and Palaeon-
tology of Queensland and New Guinea, p. 222, pi. 8, fig. J2.

Frotoretepora ampla Lonsdale, de Koninck, 1878, Mem. Soc. Hoi), dcs
Sciences de Liege, Ser. 2, Tome YIT, p. 42, t. 8, hgs. 5 a - c.

[non] Fene Stella ampla Lonsdale, 1844, in DarAvin, Geological Observations
on Volcanic Islands, p. I(i3.

Polypora iumula Laseron, 1918, Jour. Hoy. Soc. X.S.W.^ Vo\. LII, p. 191,
pi. VII, tig. 3, pi. IX.

Polypora tripliseriata Bassler, 1929, Palaontologie von Timor, Lief XVI,
XXVIIL Permian Hrvozoa of Timor, p. 79, pi. CCXLll (18), figs.
14-16.

Polypora looodsi Etheridge (Crockford), 1941, Jour. Hoy. Soc. N.S.W., p.
41-1, pi. X^’III, fig. 1, p\. XIX, tig. 1.

St)eeimens of this s])ecies occur in the material from three of the locali-
ties in the Kimberley District; the characters shown agree with those of the
neotype.

Kemarks: In Now South Wales this species is abundant in the Eene-
stella Shales (Branxton Stage of the Tapper Marine Series) in the Hunter
Kivcn* District, and in the riladulla Mudstones (probably about the same
horizon in the Upper Marine Series) of the South Coast; it F not knoAvn
from any other horizon in Ncav South Wales. In Tasmania it occurs at
Marlborough, and in the Grange Stage near Hobart; it has l)een recorded
by Etheridge from Queensland, and occurs in the Springsure District (Con-
suelo Ck., two miles above Cattle Ck. ; Reid, 1930, p. 157, locality 9). It
occurs also in the Callytharra Stage of the Wooramel R. District. Polypora
tripliseriata occurs in the Bitaoeni Beds at Kampong Apna in Timor.

Polypora multiporifera sp. nov.

Plate 3, fig. 4.

Holotype: E. ;18441, Australian Museum (’ollection (Coll. 11. Cok'v).
Horizon and locality; Calceolispongia Stage jf the Wandagee Series,
Syncline on Minilya Hiver. lialf mile AVest of Coolkilya Pool.

Polypora with five to six rows of zooecia, and from six to fourteen
zooecia to a fenestrulc ; branches slightly convex; fenestrules long, irregular
in length, aiul rather luirrow ; nodes small, fairlg abundant.

Tiiei‘e are i'rom 1-5 to 2-5 fenestndes vertically, and al)Out tiv(' branches
horizontally, in 10 mm.; t!ie branches are straight, and are rathei’ rounded
on lh(' celluliferous surface, so that the lateral roAvs of a]>erture'^ may open
towaials the fenestrulc's ; they are from about 0 -7 to 0-9 mm. Avide Avhere
there* are* four, from 0-97 to 1-1 mm. Avhe*re* thei'e are five, and about 11.

178

Joan .M. Ckockfohd.

inni. where thev(* are rows of zooeeia ; the width increases rather raiddly
before branching', which occurs at relatively short intervals. Small nodes
were appaiamtly rather abundant, though they arc; now frequently weathered
away; generally not more than two occur adjacent to each zooecial apertuia .
The whole of the celluliferous surface is hnely tuberculate; the dissepiments
>how numerous tine transverse striae where they are slightly worn. There
are usually Hve to six I'ows of zooecial ai)crtures, with from six to seven
before, and from three to tour after, branching; tin* apertures ai*e cii’culai*.
about 0.16 mm. in diamet(‘r; no peristomes are shown: the distance betweeii
the centres of successive ai)ertures is i'rom 0-J8 to 0-40 mm., and about
twenty-two apertures occur in 10 mm. with from six t<> fourteen o])posite
each fenestrule. The fenestrules are fi'om 2 \ to o (i mm. in length, and
from 0*65 to 11 mm. in width; the dissepiimaits, whi(-h are not celluli-
ferous, are from 0-46 to 0-6 mm. in width. The re\'erse surface is mtt
shown. The cells are short and are rhomboidal in shape; they are from
0 J2 to 0-38 mm. in length, and are about 0-24 mm. iii w’idth at their widest
part; the extremities of successive cells in the same row are not in conta<d.
Xo surface cells are developed.

Hemarks: The rather numerous small nodes, tin* absence' (d' any
rounded sui’face cells, and the short, broad zooeeia separate this species
from Pohf })()}'(( nKignufeucsfrafa Crockfoi'd, from the Lower and I pi)fu
Marine Series of Xew Soutli Wales.

Folypcra s]). nov. indet.

Plate 2, tig. 5.

Horizon and locality: Xooiicanbah Series; scarp two miles Last of
Christmas (’k. Homestead. (Specimen 200o2 a, Lniversity of ^^estern Aus-
tralia Collection; Coll. A. Made).

Polypora tcith about eight roivs of iooeci((, and tnfh from jiiih' b*
fiftreu apertures to a feucMruJe; nodes not developed.

The specimen is a I'ather weathered cast of the cellulii erous s\irlaee.
and the form of the colony is not well shown, though it was pi'o})ably in-
fundibuliform, with the celluliferous surface internal. Tlnere are from 3
to 4 branches horizontally, and from 1 o to 2 fenc'^tniles vertically, in
JO mm. The branches bifui-cate at relatively very fretiuent intervals, gener-
ally less than the length of one fenestrule apart ; they ai'e usually from
J-2 to l-7)5 mm. in width, but broaden considerably just before bifurcation.
There are normally about seven or eight rows of zooecial apertures; thc-
branches are thick and very convex, and the latei'al rows of apertures open
towards the fenestrules. The apertures are circular, about 0-14 mm. in dia-
meter, and the distance betAveen the centres of successive apertures is about
0-49 to 0-6 nmi.; the number of zooeeia in the length of one fenestrule
and one dissepiment A’aries with the length of the fenestrules fi'om about
9 to 15. Xo nodes are shown. The fenestrules are oval, from 3-3 to 6-9 mm.
in length, and from 0-86 to 1-41 mm. in width at their widest ]>art; the dis-
sepiments, which ex})and considerably at their junction Avith the branches, are
from 1-2 to 1-6 mm. in Avidth ; they are not celluliferous. The i-cA erse surface
is not shoAvn.

Remarks: This species most closely resembles Polifjiora yigantea M aagen
and Pichl, from the Middle Productus Limestone of the Salt Range, but is.
distinguished by its narrower fenestrules and broader dissepiments, and by
its much more frequent bifurcations.

Bryozoa from the Wandagee and Nooncanbah 8erip:s
(Permian) of Western Australia.

179

ACKNOWLEDCijVIKNTS.

1 wish to thank Dr. C. Teidiert and Dr. A. Wade for the information
whieh they ha\ e given me regarding the localities and horizons of the speci-
mens described^ and Dr. 1. A. Brown for her help in the preparation of this
paper. The majority of the specimens were lent to me by the Department
of Geology at tin' I’niversity of Western Australia, but some specimens lent
by the Australian Museum have also l)een used. Three species are recorded
from (Queensland with the permission of Shell ((Qld.) Development Pty.,
Ltd.

This work has l)een carried out during the tenui-e of a
Scholarshi[) at the rniversity of Sydney.

ScieiU'e

Kes(‘ar(ii

BIBLBKJKAPHV.

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(iravenhage.

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S[>ecial Reference to Fitzi-oy Basin, and the Possibilities of Mineral Oil,
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3. Bretnall, R. AV., 1926: Palaeontological Contributions to the Geology of

\V(Mtern Australia, Series A"I1., No. XIIL: Bes('riptions of some AVesteru
Australian Fossil Polyzoa, IF../. Geol. Surrey, Bull. 88.

4. ('hapman, P., 1904: On a Collection of Ppper Palaeozoic and Mesozoic

Fossils fiom AVest Australia and (Queensland, in the National Museum,
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7a. Crockford, .loan M., 1941: Permian Bryozoa of Eastern Australia, Part I.:
A Revision of Some Previously-named species of Fenestrelliiiidae
{ Fenestellidae') , Jour. Boy Soe. N.S.W., Ah)l. LXXIAk, Pt. IV. , pp.
397-418.

8. Flias, M. K., 1937: Stratigrai»hic Signilicance of Some Ijate Paleozoic

Fenestrate Bryozoaiis, Jovr. Paleontology, \o\. 11. , No. 4, pp. 306-334.

9. Etheridge, R., jur., 1889: Remarks on Fossils of Permo-Carboniferous Age,

from North-AVestern Australia, in the Alacleay Museum, Proc. Linnean
Society of A^..S'.JF., Ser. 1 L, A'ol. iv., ]>t. 2, pp. 199-214.

10. Etheridge, R., jnr., 1903: Palaeontological Contributions to the Geology of

AVestern Australia; L: Descriptions of Carboniferous Fossils from the
Gascoyne District, AVestern Australia, IT. A. GeoJ. Survey, Bull. 10.

11. Etheridge, R., jnr., 1907: Palaeontological ('Contributions to the Geology of

AA^estern Australia; IL: Descriptions of Carboniferous Fossils from the
Irwin River, fV.A. Geol. Survey, Bull. 27, pt. 1A\

12. Etheridge, R., jnr., 19(D a: Official Contributions to the Palaeontology of

Routli Australia, No. 19— Fossils of Port Keats Bore, Thirty Miles North
of Fossil Head, Treachery Bav, Supplement to Parliamentary Paper 55
of 1906.

13. Etheridge, R., jnr., 191-): Palaeontological Contril)utions to the Geology of

AAMstern Australia, Series A^., No. X, AAMstern Australian Carboniferous
Fossils, chielly from Mt. Alarmion, Lennard River, West Kimberley,
W.A. Geol. S^irvey, Bull. 58.

14. Foord, A. H., 1890: Notes o7i the Palaeontology of AVestern Australia,

Geol. Mag., N.S., Dec. III., A"ol. A'll., pp. 145-155.

15. Frederiks, G. N., 1915: La Faune Paleozoif|ue Superieure des Environs de

la vide Krasnooufimsk, Memoircs due Comite Gcologique \Petrograd\,
Nouvelle Serie, Livraison 109.

16. Fritz, Madeleine, 1932: Permian Bryozoa from ATmeouver Island, Boy. Soc.

Ca^iada, Transactions, Third Series, A'ol. XXAM., Section lA'., pp. 93-110.

ISO

JOAX ^1. CUOCKFOHI).

17. Hill, Dorothy, 1936: The Permian Corals of Western Australia, Jour. Hoy.

Soc. W.A.j Vol. XXIII., pp. 43-62,

18. Hinde, G. J., 1890: Kotes on the Palaeontology of Western Australia, Corals

and Polyzoa, Geol. Mag., N.S., Dec. III., Vol. VII., pp. 194-204.

19. Hosking, Lucy F. V., 1931: Fossils from the Wooramel District, Western

Australia, Jour. Hoy. Soc. W.A., Vol. XVII., pp. 7-52.

20. Hudleston, W. H., 1883: Notes on a Collection of Fossils and Eock Specimens

from West Australia, North of the Gascoyne River, Q.J.G.S., Vol.
XXXIX., pp. 582-595.

21. Maitland, A. Gibb, 1909: Geological Investigations in the Gascoyne, Ash-

burton, and West Pilbara Goldfields, IV. A. Geol. Survey, Bull. 33.

21a. Martin, R., 1931: De Palaeontologie en Stratigraphic van Nederlandsch
Oost-Indie, Brvozoa, Leidischc, Geologische Mededeelingen, Deel V.,
pp. 390-395.

22. Nikiforova, A. I., 1933: Middle Carboniferous Brvozoa of the Donetz Basin,

Trans. Vnited Geological and Prospecting Service of the U.S.S.E..
Fascicle 237.

23. Raggatt, H. G., 1936: The Geology of the North-West Basin, Western

Australia, -with Particular Reference to the Stratigraphy of the Permo-
Carboniferous, Jour. Hoy. Soc. K.S.IV., Vol. LXX., pp. 100-174.

24. Raggatt, H. G., and Fletcher, H. 0., 1937; A Contribution to the Permian-

Upper Carboniferous Problem, and an analysis of the Fauna of the
Fpper Palaeozoic (Permian) of the X’orth-West Basin, Western Aus-
tralia, Austr. Museum Hecords, Vol. XX., No. 2, pp. 150-184.

24a. Reid, d. II., 1930; Geology of the Springsure District, Qucey^sland Govt.
Mining Jour., \'ol. XXXI. (April'), pp. 149-157.

25. Teicliert, C., 1939; Recent Research Work in the Permian of Western

Australia, Austr. Jour. Soivnee, Vol. IT., No. 1, pp. 5-7.

26. Teicliert, C., 1940: Helicoprion in the Permian of Western Australia, Jour.

Paleontology, Vol. 14, No. 2, pp. 140-149.

27. Teicliert, C., 1941: Upper Palaeozoic of Western Australia; Correlation and

Palaeogeographv, Bull. Atner. Assoc. Petroleum Geologists, Vol. 25,
N'o. 3, pp. 371-415.

28. Wade, A., 1937: The Geological Succession in the West Kimberley District

of We.stern Australia, Austr. Assoc. Adv. Science, Heport, Vof. XXIII.,
pp. 93-96.

29. Wade, A., 1938: Geological Map of Part of the West Kimberley Division.

Western Australia, Preiiared on behalf of the Freney Kimberley Oil
Co.; Canberra.

PLATE I.

Figure I. Minilya princeps sp. nov.: Part of the celluliferous surface of the
holotype, showing the two rows of zooecial apertures, and the double row of
weathered nodes along the centre of the branch (No. 20945, Univ. of Western
Australia), x 10.

Figure 2. Fenestrellina disjecta sp. nov.: Part of the celluliferous surface of
the holotype (No. 2793a, Univ. of Western Australia), x 10.

Figure 3. Fenestrellina horologia Bretnall: Part of the celluliferous surface of a
specimen from the Nooncanbah Series at Keevie’s Well, 8 miles north of Mt,
Anderson Home.stead (N’o. 20942, University of Western Australia), x 10.

Figure 4. Fenestrellina valotlis sp. nov.: Part of the celluliferous surface of tlie
holotype (No. 2793c, Univ. of Western Australia), x 10,

Figure 5. Minilya duplaris sp. nov.: Part of the celluliferous surface of the
holotyjie (No. 2793d, Univ. of Western Australia), x 10.

Figure 6, Fenestrellina horologia (Bretnall) : Part of figure 3 enlarged to x 20,
to show the single row of high nodes, and the apertures, which are stellate
where they are best presein'ed.

Figure 7. Minilya duplaris sp. nov.: Part of figure 5 enlarged to x 20, to show
the arrangement of the apertures and the double row of nodes.

Bryozoa from the Wandagee and Xooncanbah Series
(Permian) of Western Axjstralia.

181

PLATE I.

182

JOAX 'M. Crockfokd.

PLATE II.

Figure 1, 2. FenesirelJina ruidacarinaia sp. nov. : Part of the celluliferous sur-
face of the holotype (Xo. 2757Uj Fniv. of Western Australia), x 10 and x 20.

Figure 3- FenestreUina columnaris sp. nov.: Part of the celluliferous surface of
the holotype (No. 20949, Univ. of Western Australia), x 10.

Figure 4. Polypora retificis sp. nov.: Part of the celluliferous surface of the
holotype (Xo. 20951, Univ. of Western Australia), x 10.

Figure 5. Polypora sp. nov.: Cast of the celluliferous surface (Xo. 20952a,
University of Western Australia — colony on the lower half of the specimen),
natural size.

Bkyozoa from the Wandaoee and Nooncanbah Series
(Permian) of Western Australia.

183

PLATE II.

18 i

JoAN' M. CliOCKI-OItlJ.

PLATE TII.

Figure 1. rojypora fovea sp. nov. : Purt of the celluliferous surface of the
holotype (No. 20947, Pniv. of Western Australia), x 10.

Figure 2. Polypora ^vo<}(]si (Ktheridge): Part of the i-elluliferous surface of a
speciineu from the Nooncanbali Series at Keevie's Well. 8 miles north of
Mt. Anderson Homestead (No. 20943, University of Western Australia), x 10,

Figure 3. Polypora fovea sp. nov.: I^’act of the celluliferous surface of a topo-
type (No. 20940, LTniversity of Western Australia), x 10.

Figure 4. Polypora multiporifcra s\). nov.: Part of the celluliferous surface of
the holotype (No. F. 38441, Australian Museum), x 10.

(Photographs by II. G. Gooch.)

15ry()zoa from the Wandagee and Nooncanbah Series 185
(Permian) of Western Austrama.

PLATK III.

A REVISIOJJ OF SOME PREVIOUSLY DESCRIBED SpECIES OF JBrYOZOA

FROM THE Upper Palaeozoic of Western Australia.

187

9.— A REVISION OF SOME PREVIOUSLY DESCRIBED
SPECIES OF BPvYOZOA FROM THE UPPER
PALAEOZOIC OP WESTERN AUSTRALIA.

By Joan M. Crockford, M.Sc.

Bead 14th April, .1942: Published 24th March, 1944.

Communicated by Professor E. de C. Clarke.

CONTENTS.

Page.

Introduction ... ... ... ... ... ... ... ... ... ... 187

Description of Species : —

Genus Fenestrellina d’Orbigny.

Fenestrellina affiuensa (Bretnall) ... ... ... ... ... ... ... 188

FenesireUina horologia (Bretnall) ... ... ... ... ... ... ... 189

Genus lAjropora Hall.

Lyro'pora erhosoides (Etheridge) ... ... ... ... ... ... ... 191

Genus Ramipora Toula.

Ramipora amhrosoifhs (Bretnall) ... ... ... ... ... ... ... 193

Acknowledgments ... ... ... ... ... ... ... ... ... ... 195

Bibliography ... ... ... ... ... ... ... ... ... ... 195

INTRODUCTION.

Four species of Bryozoa, Lyropora (?) erkosoides Etheridge, Fenestella
affluensa Bretnall, Fenestella horologia Bretnall, and Aetomocladia amhrosoides
Bretnall, are hei’o revised from the type material in the Western Australian
Geological Survey Collection. In the original descriptions of these species
the number of the holotype of eacli species is quoted, but in labelling the
specimens the same number was used for all the specimens referred to the
species, and in some cases the original labels are now missing ; lectotypes
are therefore chosen from the syntypes of the species revised, and the speci-
mens liave been recatalogued.

DESCRIPTION OF SPECIES.

Phylum BRYOZOA Ehrenberg.

Class GYMNOLAEMATA Allman.

Order CRYPTOSTOMATA Vino.

Faiuily FENESTRELLINIDAE Bassler.

Genus FENESTRELLINA* d’Orbigny.

Feyiestrellina d'Orbigny, 1849, Revue ct Magasin de Zoologie, 2e Sdr.,
Tome I., p. 501.

Homonym •

Fenestella Lonsdale, 1839, preoccupied foj- a peleeypod, FenesteUa
Bolton, 1798.

Genotype : Fenestella crassa McCoy, 1845.

* An application for saspension of the Rules of Zoological Nomenclature for the generic name Fenestella
Lonsdale 1839 has been submitted to the International Commission on Zoological Nomenclature by G. E.
Condra and M. K. Elias {Journal of Paleontology, Vol. 15, no. 4, pp. 565-56H).

k 44/44

188

Joan M, Crockforb, M.Sc.

Infundibuliform or flabellate Fenestrellinidae, with two rows of alternating
zooecia on the branches, except before bifurcation, when the number of rows is
increased ; rows of zooecia separated by a median carinn, plain or more fre-
quently with a single row of nodes {acantho pores) ; dissephnents not cellnliferous;
internal structure and reverse surface as for the family.

Fenestrellina affluensa (Bretnall).

(Plate I., fig. 6.)

Fenestella affluensa Bretnall, 1926, W.A. Geol. Survey, Bull. 88, p. 16,
pi. I., fig. 8.

Fenestella affluensa Bretnall, Hosking, 1931, Jour. Roy. Soc. W.A., Vol.
XVII., p. 12.

Lectotype : Specimen 2 /2405E, Western Australian Geological Survey

Collection ; on specimen figured by Bretnall, 1926, pi. III.

Horizon and locality ; Between the top of the Lyons Series and the top
of the Byro Series, Gascoyne River District, W.A.

Fenestrellina with three to four zooecia to a fenestride ; carina slight,
rounded ; nodes large.

The form of the colony is not shown ; the lectotype is about Do cm.
long and D1 cm. wide. There are from 12 to 14 branches horizontally, and
8 fenestrules vertically, in 10 mm. ; the branches are straight, normally from
about 0'4 to 0*53 mm. in width, but rather narrower — about 0*35 mm. —
where they are slightly weathered ; on the celluliferous surface the carina is
poorly developed, being represented only by a I’ounded area free from ceils
along the centre of each branch, sej)arating the two rows of alternating cell
apertures. Nodes are wdl developed, but are only shown where the surface
is best preserved ; they arc very large and blunt, and are slightly elongated
parallel to the length of the branch ; theii’ width at the top is up to 0*33 mm.,
and the distance between the centres of successive nodes is from 0*68 to
0*86 mm. The apertures are normally circidar, about 0*14 mm., in diameter,
but they may be much enlarged by weathering ; they do not generally project
into the fenestrules. Thin, raised peristomes are shown where the sui-face
is well-preserved. The distance between the centres of successive apertures
is from 0*29 to 0*37 mm., and about thirty occair in 10 mm. The fenestrules
are oval — the dissepiments expand considerably at their junction with the
branches — and are from 0*85 to 1*2 mm. in length and from 0*3 to 0*5 mm.
wide ; the width of the dissepiments is from 0*25 to 0*36 mm. Bifurcation
occurs at rather distant intervals, and increase to three rows of zooecia occiirs
just before branching. The thickness of the branches is about 0*65 mm.

In specimens from the same locality, but not sjmtypes, the reverse surface
shows a number of fine longitudinal striae where it is slightly weathered ;
the dissepiments are very nearly as thick as the branches, and both are evenly
rounded.

Remarks : There are three specimens labelled “ 17 ” (the number given
by Bretnall as that of the holotype) in the collection ; of these the largest
and best preserved is here chosen as the lectotype, and the measurements
given in the above description were taken on it ; there is a second, very small
specimen on the same piece of shale, and a third, very much weathered speci-
men on a piece of limestone from the Callytharra Stage at Fossil Hill, Wyndham
River. Both of these show fenestrules slightly longer than the average shown

A EEVISION OF SOME PREVIOUSLY DESCRIBED SpECIES OF BrYOZOA 189
FROM THE Upper Palaeozoic of Western Australia.

by the lectotype, though they fall within the range of variation of it ; they
are weathered so that the nodes are not visible, but appear to be conspecific
with the lectotype.

Fcnestrellina affiue)isa is distinguished from associated species by its
thick branches, and large, well spaced nodes.

Fenestrellina horologia (Bretnall).

(Plate T., fig. .] ; Plate II., fig. A.)

Fe-nesfella horologia Bretnall, 1926, W.A. Geol. Surv&y, Bull. 88, p. 15,
pi. I., fig. 6.

[non] Feyiestella horologia Bretnall, Hosking, 1931, Joui\ Roy. Soc, W.A.,
Vol. XVII., p. 13, pi. IV., fig. 3.

Fenestrellina horologia (Bretnall), Crockford, 1944, Jour. Roy, Soc. W.A.,
Vol. XXVIII., p. 167, pi. I., figs. 3, 6.

Fenestella parviuscula Bassler, 1929, Paldontologie von Timor, Lief. XVI.,
XXVIII., p. 76, pi. 17 (CCXLI.), figs. 8-13.

Fenestella parviuscula Bassler, Martin, 1931, Ue Palaeontologie en
Stratigraphie van Nederlandsch Oost-Indie, Bryozoa, p. 391.

Fenestella parviuscula Bassler, Fritz, 1932, Roy. Soc. Canada, Trans-
actions, Third Series, Vol. XXVI., Sect. IV., p. 99.

Fenestrellina parviuscula (Bassler), Elias, 1937, Jour. Paleontology. Vol.
11, No. 4, p. 314.

Neotype (? Lectotype); Specimen 2/2405C, Western Australian Geo-
logical Survey Collection ; on specimen figured by Bretnall, 1926, pi. III.

Horizon and locality : Between the top of the Lyons Series and the top
of the Byro Series, Gascoyne River District, W.A.

Fenestrellina with two zooecia to a fenestrule ; cariyia low ; nodes sharp,
relatively high, about two to a fenestrule ; fenesfrules hoyir-glass shaped.

The foi’m of the colony is not shown ; there are about 18 branches hori-
zontally, and from 16 to 18 fenestmles vertically, in 10 mm. The branches
arc straight, from 0-31 to 0*37 mm. in w'idth, and show two rows of regularly
alternating zooecial apertures, separated by a slight median carina. Bifurca-
tion occurs at rather distant intervals, and increase to three rows of zooecia
occurs just before branching. The apertures are circular, from 0*08 to O-l
mm. in diameter, and are placed on the slightly sloping sides of the branches ;
the peristomes are not well shown. The apertures are generally very regularly
placed, with one opposite the centre of each fenestrule and one at the end
of each dissepiment ; the former project into the fenestrules — though the
apertures open upwards and not into the fenestrules — and give them their
characteristic hour-glass shape. The distance between the centres of succes-
sive apertures is from 0*24 to 0*3 mm., and about thirty-seven occur in 10
mm. The nodes are small and sharp, and arc placed in a single row on the
carina ; they are from 0*24 to 0-3 mm. apart. The fenestrules are from 0*32
to 0-51 mm. in length, and from 0*22 to 0-33 mm. — generally about 0*25
mm.— in width ; the dissepiments are from 0-11 to 0* ] 7 mm. in -width. The
reverse surface is not shown.

190

Joan M. Crockford, M.Sc.

Remarks : Fenestrellina horologia w’as described by Bretnall from material
from the Gascoyne River District ; tlie holotype was given as “ Geological
Survey of Western Australia (specimen 16).” Miss Hosking (1931, p. 13)
stated that “ the holotype of F. horologia (specimen 16 on 10930) is not to b(‘
found in the Geological Survey Collection” but (footnote) ‘'A crumpled
fragment of a Fencstella zoarium is labelled Holotype, but the distinguishing
number and the greater part of the zoarium have been broken off.” Speci-
mens which she identifiecl as Fenestella horologia from the Wooramel River
District showed a double row of nodes on the carina ; she states that the
specimen in the Geological Survey Collection was much the same as these,
but was “* too crumpled for accurate comparison,” and does not say whether
a single or a double row of nodes was sho^Mi.

There is at present no specimen in this collection labelled ” 16,” and
no specimen labelled “ Holotype.” The labels of many of the specimens
have been wholly or partially destroyed, and there are two eonspecific frag-
ments of Fenestrellma, whose labels have been almost entirely removed, on
a specimen of calcareous shale from the Gascoyne River District, figured by
Bretnall on Plate III. (this specimen shows several more labels now than at
the time is was photographed). Both of these show a slight carina between
the two rows of apertures, and over the best preserved parts of the zoaria
this carina shows a single row of small, high nodes ; both show \-ery well
the hour-glass shape of the fenestrules described by Bretnall ; it seems
probable that these were the specimens used for the original description,
and one of them is here chosen as the neotype (? lectotype). A single specimen
of a second species showing only two zooecia to each fenestrule is present
on the same piece of shale ; this specimen shows, very indistinctly, two rows
of small nodes on the carina, and is probably the species identified as
Fenestella horologia from the Wooramel River, described in a separate paper
as Minilya duplaris gen. et sp. nov. (Crockford, 1944, p. 173) ; this, the only
specimen of this species present in the collection, is very poorly preserved ;
it does not show any projection of the zooecial apertures into the fenestrules,
and the hour-glass shape of the fenestrules is not shown at all in this specimen,
though it is shown in specimens from other localities ; it is not possible that
Bretnall’s description could refer to this specimen.

Fenestrellina horologia appears to be the same as Fenestrellina -parviuscida
(13assler), from the Bitaoeni and Basleo Beds of Timor ; in the original de-
scription of F. parvniscula the width of the branches is given as “ about 0-2
mm.”, but the branches of the specimens figured appear to be about 0*3 mm.
wide, as in the specimens from Western Australia. Elias (1937) has tlealt
^vith the relationships of this and similar species. F. parviuscida has been
recordc'd from the Permian of Vancouver Island, llie fenestrate part of
Lyropora erkosoides (Etheridge) is very similar in size, but is differentiated
by its more widely spaced nodes. Fenestrellina pectinis (Moore), which has
been recorded by Chapman from the Callytharra Stage at Callytharra Springs,
is similar iu appearance, but is a smaller species with more closely sjiac^ed
nodes.

Genus LYROPORA Hiili.

Lyropora Hall, 1857, Proc. Amer. Assoc. Adv. Sci., Vol. X., p. 179.

Lyropora Hall, Ulrich, 1890, Geol. Surv. Illinois, VIII., pt. II., Sect. IV.,
pp. 396, 580.

Lyropora Hall, Nickles, and Bassler, 1900, U.S. Geol. Surv., Bidl. 173,
pp. 39. 309.

A REVISION OF SOME PREVIOUSLY DESCRIBED SpECIES OF BRYOZOA 191
FROM THE Upper Palaeozoic of Western Australia.

Lyropora Hall, McNair, 1937, Michigan University Museum of Paleojitol-
ogy Contributions, Vol. V., No. 9, p. 114.

Genotvpe (selected bv Ulrich, 1890) : L'tiropora subquadravs (Hall),

1857 (L. lyra (Hall), 1857).

Synonyms : —

Dictyoretmon Whitfield, 1904.

Lyroporella Simpson, 1895.

Lyroporina Simpson, 1897.

Zoariuni fenestrate ; colony flahellif or m, with the sides near the base strongly
thickened to form the lateral supports ; brayiches with two or more rows of zooecia,
dissepimemts without zooecia ; reverse swface and internal structure as in Fenos-
trellina.

Lyropora erkosoides (Etheridge).

(Plate I., fig. 2 ; plate II., fig. B.)

Lyropora (^) erkosoides Etheridge, in Bretnall, 1920, W.A. Geol. Survey,
Bull, 88, p. 11, pi. II., figs. 1, 4.

Lectotype : Specimen 2 /2404, Western Australian Geological Survey

Collection; figured by Bretnall, 1926, pi. II., fig. 1.

Horizon and locality ; Callytharra Stage (?), Well A 20, Daiirie Creek,
Gascoyne River District, Western Australia.

Lyropora with two rows of zooecia, and two zooecia to a fenestride ; carina
slight . nodes sharp, high, evenly spaced.

The zoarium is flabellate ; the base is small and pedunculate, and the
sides of the colony above the base are strongly thickened, and form the lateral
supports characteristic of the genus. These lateral supports diverge at an
angle of about 70°, and the zoarium is celluliferous on the convex surface ;
the holotype reaches a height of 4 cm. above the base, and a width, at the
top, of about 4 cm. There are about 22 branches horizontally, and 18 fenes-
trules vertically, in 10 mm, The branches are straight, and are usually from
0*22 to 0*29 mm. in width, but they become much broader towards the sup-
ports, and before bifurcation. There are two rows of zooecial apertures,
separated by a very slight median carina on which there is a single row of
high nodes, rather elongated along the carina at their bases. The nodes are
placed from 0*3 to 0*37 mm. apart. The apertures are circular, about
0*09 mm. in diametei' ; slight peristomes arc shown — these are best developed
on the side towards the fenestrules. There are two apertures to a fenestrule,
and over part of the specimen these are arranged so that one is opposite the
end of each dissepiment, and one at the centre of each fenestrule, giving the
fenestrules an hour-glass shape ; over most of the specimen, however, the
arrangement is not regular, and the fenestrules are oval in shape. The dis-
tance between the centres of successive apertures is from 0-24 to 0-32 mm.,
and about thirty-seven occur in 10 mm. The fenestrules are from 0-35 to
0-46 mm. in length, and from 0*17 to 0-29 mm. — generally about 0*2 mm. —
in width ; the width of the dissepiments is from 0-13 to 0-21 mm. On the
reverse surface both branches and dissepiments are evenly roimded, and
they are of about the same thickness — their thickness varies, however, with
the distance from the lateral supports. The outer layer of the reverse surface
is finely granular ; within this a few fine longitudinal striae are shown. Bi-
furcation occurs at very distant intervals, and increase to three rows of zooecia
occurs just before branching.

192

Joan M. Crockford, M.Sc.

Remarks : There are two specimens labelled “ 13 " (the number given

by Rretnall for the holotype of this species) in the material described by
Bretnal!. The above description is of the figured specimen, here chosen as
lectotype. A second speciimm, from the Callytharra Stage at Fossil Hill,
Wyndham River, does not appear to belong to the same species as tlie lectotype,
from which it differs in the shape of the lateral supports ; only a \’ery small
portion of the fenestrate part of the /,oarium is preserved, along the sides
of the lateral supports, and tiiis is decidedly finer than the fenestrate part of
the lectot;v 7 :)e.

Ktheridge referred this species teiitatively to the genus Lyropora Hall,
but, as the celluliferous surface of the specimen was not exposed, considered
it possible that the species might belong to one of the genera proposed by
Simpson for forms diflering from the type of Lyropora in the number of rows
of zooecia normally present on the branches. The type of Lyropora, Lyropora
fiiibquadrans (Hall) (L. lyra (Hall)), selected by Ulricli in 1890, shows from four
to five rows of zooecia on the branches ; Lyroporidra Simpson, 1897, is a
synonym of Lyropora, since it possesses the same genotype. Lyroporella
Simpson, 1895, was proposed for forms with two rows of zooecia, the genotype
being Lyropora quincuncialis (Hall) ; Nickles and Bassler (1900, p. 310), who
considered that the only character of generic importance is the development
of lateral supports, and McNair (1937, p. 1 14) consider Lyroporella a synonym
of Lyropora.

(family SULCORETEPORIDAF Bassler.

Gsnus RAMIPORA Toiila.

Ramipora Toula, 1875, Neues Jahrbuch fur Mineralogie, 1875, p. 230,
pi. X., figs. 1, la.

Ramipora Toula, Shulga-Nesterenko, 1933, United Geological and Pros-
pecting Service of the U.S.S.R., Trudi, Fasc. 259, pp. 32, 54.

Genotype : Ramipora hochstettej'i Toula, 1875.

Synonym : —

Aetomocladia Bretnall, 1920.

Genotype: Aetomoedadia a)}ibrosoides Bretnall, 1920.

Zoarium pinnate, ivith the lateral branches joined by a third- set of branches,
which may themselves join to foryyt a further set of branches ; branches bifoliate,
dividing in a plane at right angles to the mesial lamina ; mesial lamina running
from the centre of the obverse (poriferous) surface to the centre of the re-verse (non-
poriferous) surface, and projecting slightly above these surfaces so that both are
carinate ; zooecia sub-tubular, without hemisepta, arranged in more or less regidar
rows on each side of the carina of the poriferous surface, but not separated intern-
ally by vertical plates ; lunaria absent or poorly developed. ; diaphragms extremely
rare ; acanthopores and mesopo7'es absent ; vesicular tissue well-developed in-
ternally, interapertural spaces solid at the surface.

Ramipora differs from Goniocladia Ftheridge in the form of the colony,
which is reticulate in Goniocladia. Sulcoretepora d’Orbigny (Cystodictya Ulrich)
differs in its mode of branching (in the plane of the mesial lamina instead of at
right angles to it), and in the form of the zoarium, and internally in the presence
of vertical double plates between the rows of cells.

A REVISION or SOME Previously Described Species of Bryozoa 193
FROM THE Upper Palaeozoic of Western Australia.

I^retnall (1926, p. 31) (lf;scrib('(l Aeiomocladia as a new genus “related
to Finnatopora Vines” the genotype b(‘ing Aetom,ocladia ambrosoides Bretnall,
from tlie Gascoyne River Distri(^t, W.A., and Fossil Hill, Wyndhain River,
W.A. Bretnall did not study tlu^ inbTnal structure of tlu^ g(‘noty[)e, and
it has therefore been considered a member of the AcanthocladiidatL The
internal structure, as is shown under the description of the species given
below, is the same as that described by Bassler and Moore for species of Gonio-
(iladia, and by Shulga-Nesterenko for Goniocladia and Ramipora, and indicates
that Aetomocladia is a member of the Sulcoreteporidae allied to these genera ;
the form of the colony separates Aetomocladia from Goniocladia., and indicates
identity with Ramipora (sens. str.).

Ramipora ambrosoides nh''tnall).

(Plate I., figs. 3-5 ; plate, IT., figs. C-F.)

Aetomocladia, ambrosoides Ihvtnall, 1926, W.A. Geol. H'urvey, Bull. 88,
p. 21, pi. I., fig. 4.

Aetomocladia ambrosoides Bretnall, Hosking, 1931, dour. Roy. Foe. W.A.y
Vol. XVII. , p. 12, pi, IV., figs. 5-6.

Aetomocladia a'tubrosoides Bretnall, Chapman, in Baggatt, Jour. Roif.
Hoc. 7V..S.W., Vol. LXX., pp. 106, 128, 148,

Aetomocladia a')uhrosoid,es Bretnall, Raggatt and Fletcher, 1937, Australian
Museum Records, Vol. XX., No. 2, p. 173.

Acanthocladia acuticostata Bassler, 1929, Paldontoloyie von Thnor, Lief.
XVI., XXVIll., p. 85, pi. 20 (CCXLIV.), fig. 13.

Lectotype : Specimen 2 /2405B, Western Australian Geologi(^al Sur\-(‘y
Collection; on specimen figuixul by lh’(‘tnall, 1926, ])1. 111.

Horizon and locality : Bedween th(> top of the Lyons Series and the top
of the Byro Series, Gascoyne Riv('r District, W.A.

Fine Ramipora, with from three to jive rows e)f zooecial apertures on each
side of the inesial lamina.

Since the lectotype does not show the form of the colony very well, para-
types from the same locality, and from tlie ('allytharra Stage at Fossil Hill,
W^mdham Rivei-, and a number of specimens from the type locality of the
('allytharra Stage in the Wooramel River District, and from the Nooncanbah
Series in the Kimb(U‘l(‘y District have been used for this description ; the
measiiD'inents given, liowever, were taken only on the lectotyp(^ and otlior
specimens on the same piece of shale.

The bas(' of the zoarium is not shown on any of the specimens ; the
zoarium is pinnate ; the branches are bifoliate, and division of the branches
takes place in a piano at right angles to the mesial lamina. Lateral (scicondary)
branches are given off from the main stem, and thc'inselves give off tertiary
branches which usually j^ass straight from one secondary bi-anch to another,
but which rarely pass outwards obliquely and join to form a further set of
branches. Subsidiai-y branches appear generally to originate at th(‘ same
level on opposite sides of a branch, but actually one is nearly always given
off very slightly before the otlier, as is shown by the junction of tlie mesial
laminae of the lateral branches with that of the original branch ; very rarely
the lateral branches on opposite sides are more widely separated. The dis-
tance between the origins of two suc(;essive pairs of latei-al branches is from

l94

Joan M. Crockford, M.Sc.

2 ■ 5 to 5-5 mm. in these specimens, and the angle at which the lateral branches
are given off is from 50° to 77° — in specimens from the Nooncanbah Series
this angle is up to 90°. The main stems are usually slightly broader and
thicker than the lateral branches, their width being from 1 • 1 to 1-5 mm., and
their thickness about 1 • 5 mm. ; the width of the secondary branches is from
0*8 to 1*4 mm., and the tertiary branches are slightly narrower.

The branches are bifoliate, with a mesial lamina whicli runs from the
centre of the reverse to the centre of the poriferous surface, and forms a slight
ridge along the centre of each surface, so that both are carinate ; the porifer-
ous surface, however, is sharply convex, with the sides sloping steeply out-
wards, while the reverse surface is rather rounded. On each side of the carina
on the poriferous surface there are three or four, less oft cm five, rows of zooecial
apertures ; the arrangement of the apertures, however, is not always regular.
The apertures are raised above the surface by the development of high peri-
stomes, particularly on the lower side of the aperture ; from sections it appears
that no lunaria are developed ; the peristomes are very (juickly removed by
weathering. When the apertures are arranged in definite rows the amount of
alternation between the apertures of adjacent rows is very variable, and often
very slight. The apertures are circular, or pyriform where they are weathered,
and are from 0- 14 to 0- 18 mm. in diameter ; they are rather irregularly spaced
on all the branches, and tlie distance between tlio ctmtres of successive apcudures
in the same row is from 0-38 to 0*76 mm. (average 0-50 mm.), and about
eighteen apertures occur in 10mm. No acanthopores are developed; the
surface between the apertures is solid. The reverse (non-poriferous) surface
is finely granular.

Internally the zooecia are sub-tubular, and the length of individual zooecia
is about 1 mm. Diaphragms appear to have been very rarely developed near
the proximal ends of the zooecia. The median tubules of the mesial lamina are
well-showm, and vesicular tissue is well-developed, particularly near the reverse
surface, and at the origin of subsidiary branches.

Remarks : In the original description of the species the genotype ”

is given as “ specimen 12,'’ but there are seven specimens labelled 12 ” in the
collection ; these belong to three species, but one specimen, belonging to
the species to whicli the original description most closely applies, is labelled
Genotype,” and is hero chosen as the lectotype of the spe'cies.

Of described species of Ramvpora, the genotyjie, R. hochstetteri Toula, and
a specimen recorded by Reed (1925, p. 107, pL X., figs. 9, 9a), as R. cf. hoch-
stetteri from India, most closely resemble this species ; they differ, however,
in tlieir broader branches and larger measurements. Ramipora uralica Stucken-
berg, from the Upper C’arboniferous of the Urals, appears to have had lateral
branches placed about the same distance apart, but the branches are much
broader.

The type specimen of Acanthocladia acuticostaia Bassler, from the Basleo
beds of Timor, appears to be a specimen of this species.

A specimen described as Ramipora sp. by Etheridge (1907, p, 14) from
the Port Keats Bore appears to have differed from this species in the form
of the zoarium and in the much closer spacing of its lateral branches.

A KEVISION OF SOME PREVIOUSLY DESCRIBED SPECIES OF BbYOZOA 195
FROM THE Upper Palaeozoic of Western Australia.

ACKNOWLEDGMENTS.

I wish to thank the Government Geologist of Western Australia for tlu'
loan of specimens from the Western Australian Geological Survey Collection
and Miss Crespin for specimens from the Commonwealth Palaeontological
Collection ; and Dr. I, A. Ihuvn for the help which she has given me with
the preparation of this paper. This work has been carried out during the
tenure of a Science Research Scholarship at the University of Sydney.

BIBLIOGRAPHY.

1. Bassler, R. S., 1929 : The Permian Bryozoa of Timor, Paldoniologie von Tiuiov, X\’l.

Lieferung, XXVIII.

2. Bretnall, R. W., 1926 : Palaeontological Contributions to the Geology of \\’est('rn

Australia, Series VII., No. XIII., Descriptions of some Western Australian Fossil
Polyzoa, W.A. Geol. Survey, Bull. 88.

.3. Crockford, Joan M., 1944 : Bryozoa from the Wandagee and Nooncanbah Seri(‘s

(Permian) of Western Australia, Part I., Jour. Roy. Soc. W.A., Vo\. XVIIh, p]>.
165-185.

4. Elias, M. K., 1937 : Stratigraphic Significance of Some Late Paleozoic Fenestrate

Bryozoans, Jour. Paleontology, Vol. 11, No. 4, pp. 306-334.

5. Etheridge, R., jnr., 1873 : Description of Carinella, a New Genus of Carboniterous

Polyzoa, Qeol. Mag., Vol. X., pp. 433-434.

6. Etheridge, R., Jnr., 1876 : Carboniferous and Post-Tertiary Polvzoa, Oeol.

N. S., Dec. II., Vol. III., pp. 522-523.

7. Etheridge, R., jnr., 1907 : Official Contributions to the Palaeontology of South Aus-

tralia, No. 19. — Fossils of the Port Keats Bore, Thirty Miles North of Fossil Head,
Treachery Bay, Supplement to Parliamentary Paper No. 55 of 1906.

8. F]’itz, Madeleine A., 1932 : Permian Bryozoa from Vancouver Island, Roy. Soc.

Canada, Transactions, Third Series, Vol. XXVI., Sect. IV., pp. 93-110.

9. Hall, J., 1857 ; Observations on the Genus Archimedes or Fenestella, Proc. Amcr.

Assoc. Adv. Sci., Vol. X., pp. 176-180.

10. Hosking, Lucy F. V., 1931 : Fossils from the Wooraiiiel District, IVestern Australia.

Jour. Roy. Soc. W.A., Vol. XVII., pp. 7-52.

11. Lonsdale, W., 1839 : In Murchison, Silurian System : John Murray, London. 1S39.

12. McCoy, F., 1845 ; Synopsis of the (’arboniferous Limestone Fossils of Ireland : Tni-

versity Press, Dublin, 1845.

13. Martin, R., 1931 : De Palaeontologie en Stratigraphic van Ncdeiiandsch Oost-Indie,

Bryozoa, Leidsche Geologische Mededeelingen, Deel \^, pp. 390-395.

14. Moore, R. C., 1929 : A Bryozoan Faunule from the Upper Graham Formation, Penn-

sylvanian, of North-Central Texas, Jour. Palaeontology, Vol. 3, No. 1, pp. 1-27.

15. Nickles, J. S., and Bassler, R. S., 1900 : A Synopsis of American Fossil Bryozoa,

including Bibliography and Synonomy, U.S. Geol. Surrey, Bull. 173.

16. Orbigny, A. d’, 1849 : Sur Quelques Genres Nouveaux de Mollusques Bryozoaires,

Revue et Magasin de Zoologie, 2e Series, Tome I., pp. 499-504.

17. Raggatt, H., 1936 : The Geology of the North-West Basin, Western Australia, with

Special Reference to the Stratigraphv of the Permo-Carboniferous, Jour. Roy.
Soc. N.S.W., Vol. LXX., pp. 100-176.

18. Raggatt, H., and Fletcher, H. ()., 1937 : A Contribution to the Permian-Upper Car-

boniferous Problem, and an analysis of the Fauna of the Upper Palaeozoic
(Permian) of the North-West Basin, 'W^estern Australia, Records of the Ausfraliayi
Museum, Vol. XX., No. 2.

19. Reed, F. R. Cowper, 1925 : Pepper Carboniferous Fossils from Chitral and the Pamirs,

Pal. Indica, N. S., Vol. \T., Memoir No. 4.

20. Shulga-Nesterenko, M. J., 1933 : Bryozoa from the Coal-bearing and Subjacent Stories

of Pechora Land : Goniocladia Etheridge and Ramipora Toula, Carboniferous

and Permian Representatives of the Family Cystodictyonidae, Transactions of
the United Geological and Prospecting Service of the U.S.S.R., Fasc. 259.

21. Simpson, G. B., 1895: A Discussion of the Different Genera of the Fenestellidae,

Thirteenth Annual Report of the State Geologist [of New York] for the year
1893, Albany, 1894 [1895], pp. (>87-787. [Forty-seventh Annual Report of the
New York State Museimi of Natural Jlistory, Albany, 1894 [1895], pp. 8S1-921J.

190

JoAjsr M. Crockford, M.Sc.

2'2. yimpson, G. B., 1897 : A Handbook of the Genera of the North American Palaeozoic
Bryozoa : with an introduction on the structure of living species, Fourteenth

Annual Report of the State Geologist [of New York] for the year 1894, Albany,
1895 [1897], pp. 407-608. [Forty-eighth Annual Report of the New York State
Museutn of Natural History, Albany, 1895 [1897], pp. 407-608].

2'2. Stuckenberg, A., 1895 : Korallen und Bryozoen der Steinkohlenablagerungen des

Tral und des Timan. Mem. du Comite Geologique [Russia], Vol. X., No. 3.

24. Toula, F., 1875 : Permo-Carbon- Fossilien von der Westkuste von Spitzbergen, Leon-

hard und Bronn's Neues Jahrbuch fur Mineralogie, 1875, pp. 225-264,

25. Ulrich, E. ()., 1890 : Palaeozoic Bryozoa, Geological and Natural History Survey of

Illinois, VIIT., Pt. II., Sect. IV.

26. Whitfield, R. P., 1904 : Notice of a New Genus and Species of Carboniferous Bryozoa,

Bull. American Miiseum of Natural History, Vol. XX., p. 469.

PLATE I.

Figure 1. Fenestrellina horologia (Bretnall) : Celluliferous surface of the* neo-
type (? lectotype), x 10.

Figure 2. Lyropora crkosoides (Etheridge) : Lectotype, natural size.

Figure 3. Ramipora amhrosoides (Bretnall) : Reverse surface (Specimen 118,
Commonwealth Palaeontological Collection) from the Callytliarra Stage, west
of Callytliarra Springs, Wooramel River), x 5.

Figure 4. Ramipora amhrosoides (Bretnall) : Celluliferous surface of the lecto-
type, X .1.

Figure 5. Ramipora amhrosoides (Bretnall): Celluliferous surface (Specimen
2793 e, University of Western Australia Collection, from the Nooncanbali
Series 6V1* miles north of Mt. Anderson, West Kimberley District), x 5.

Figure 6. Fenestrellina affiiiensa (Bretnall): Celluliferous surface of the lecto-
type, X U).

(Photographs by H. G. Gooch.)

A REVISION OF SOME PREVIOUSLY DESCRIBED 8pF]C 1KS OF JiRYOZOA 197
FROM THE Upper Palaeozoic of Western Austrat.ia.

198

Joan M. Crockford, M.Sc.

PLATE II.

Figure A. FnicsfreJUna horologia (Rretnall) : Celluliferoas surface of a
topotype, X 20.

Figure B. Lyropora elcos-oides (Etheridge) : A thin section of the part of
the lectotype, x 20. (Slide (5277, Western Australian Geological Survey Collec-
tion.)

Figure C. iRa}nipom amhrosoides (Bretnall) : Longitudinal seetioiij cut loose
to the reverse surface (Slide 2793g, University of Western Australia Collection,
from a specimen from the Nooncanbah Series" tiVo miles north of Mt. Anderson,
AVest Kimberley District), x 20,

Figure D. Famipora amhroffoides (Bretnall): Longitudinal section of a
topotype,' cut rather closer to the obverse surface than C. (Slide 6278, Western
Australian Geological Survey Collection), x 20.

Figure E, F. Famipora amhrosoides (Bretnall): A^ertical sections (Slides
2793 e, f. University of Western Australia Collection, from specimens from the
Nooncanbah Series 6b', miles north of Mt. Anderson, West Kimberlev District),

X 20.

{Camera liccida diagrams.)

A REVISIOK OF SOME PREVIOUSLY DESCRIBED SpECIES OF BrYOZOA 199
FROM THE Upper Palaeozoic of Western Australia.

t'ossiL Plants from Ginoin, W.A.

20 1

](). P08SIL PLANTS FROM GINGIN, W.A.

l^Y A. H. Walkom, D.Sc.

O'Ih' Australian Musoimi, Sydnoy.*

(Platf's i.-n.)

K(‘a<l I4tli Aj)i'il. 1042; publislicd 20th March. 1044.

Abstract.

Fossil ])lants described from (Jingin indicate a Jurassic Age for beds of
clayey sandstone underlying the Cretaceous greensands. 'Plie collection in-
(dudes specimens of leaves and sporangia rc'fem'd to Isoelites^ tVie first record
of tins genus for Australia.

'fh(' (^ollc(dion of fossil ])lants her(> described was forwardt'd to nn^ by
Pioh'ssor K. de ('. (Jarke in 1037. Jt is n‘gr(‘tte<l that cinrmnstaiu^t's have
so long delayed the c,om])letion of my investigations.

The s])ecinu‘ns were obtained from brownish, fin(‘-graine<l, clay(‘y sand-
stoiu's undtu-lying the (’retaceous gn'cnsands at Oingin, W.A., and tlu^ list
of .s])ecies k'jids su]>port to Piofessor (Jarke’s statement, in a lettc'r, that he had
“fairly good (‘videnee that the chalk and gr(H*nsand lie on an erod<‘d surfact*
of th«^ plant bi*ds.*‘

'fh(‘ s])(‘ch‘S idi'iititied ari' : -

Isoetites elegans, sp. uo\'.

Clado'phlehis australis (Morris).

77/ ?’a nfvld ia fa Ibragarensls Will ko m .

'l'acnio})ti'ris spaUdata Mcdcdland.

/dilophylluni jx’ctcH (.Philli])s).

Hlatocladus plana { Fi'istnmntel).

TIkw suggest a Jurassic age for tii(‘ beds in whicli they occur, and show
clos(‘St similai'ities with th<‘ .Jurassic flora of th(^ Talbragar HimIs in N(‘w South
VVal<‘s.

Perhaps tlu> most interesting fi'ature is the presem^i* of numei‘oiis sj)eci-
UK'iis T-eferi'c'd to Isoetites. Fossil repres(‘ntatives of this group ai’t^ few in
number, and noiu‘ a})])ear to liave b('(‘n described from rocks of Jurassic agi*.
so the sp<‘cies I. elegans inaA' b(‘ the earliest sp(‘cies yt‘t known, as w(*ll as bt'ing
the first fossil s]K‘cies known fi’om Australia. genus Jsoetes has a wide

rang(‘ at tlu^ pn‘S<‘nt day b\d only six s]>ecies ap])t‘ar to be re(U)?’ded from
Australia (thr<‘<‘ from Tasmania, and one iwh from Qu('(‘nsla-nd. South Aus-
tralia, and \\7'st(‘i'n Australia).

Professor (Marke has sugg('st(‘d that tlu'se ]>lant-b(‘aring IkhIs at (lingin
may b(‘ tlu' (continuation of ])lant beds at Hullsbrook, some 30 inih^s north of
Perth. In 1031 1 examiiu‘d for him a collection of fragmentai-y jdant remains
from tlu“ Hullsbrook IhmIs and determiiu‘d tin* following .sjk*cu‘s -CVodo-
phlebis austmUs. ‘t rhyllopteris sj ThinnJeUlia sp., Taeniopteris elongata^
Nllsso}\ia sp., and Klatocladus cf. platut. This small {collection was not v(‘ry
satisfactory foi‘ (Udi'i’inination of the age of the b('ds and at the time I suggest(‘d
that possibly a Ja)W(m- Cndaceous age was indicated. As a whok* the Hulls-
brook colU’ction do('s not show (^los(‘ aihnitu^s with that from Oingin, and may
ro|)r(\s(‘nt a sonu'what Jiighei’ hori/.on.

* Publishi'd by permission of the Trustei's of t he Austi-aiian Mus(Mim.

202

A. B. Walkom, D.Sc.

LYCOPODIALES.

Isoetites elegans, sp. nov.

(Plate I.)

Leaves numeroxis, long (up to 12 cm. or more), straight or slightly curved,
about 3-4 mm, wide at base, tapering to apex, with very fine parallel striations.
Sporangia usually 1*5-2 cm. long, by 7-8 mm. Megaspores spherical, about
0*5 mm. in diameter, smooth, with equatorial ridge, and tri-radiate marking
on one hemisphere, 40-55 megaspores in a sporangium.

In the specimen figured (Plate I., fig. 1) there are some 30 leaves visible ;
in addition the bases of a number of other leaves can be observed on the vertical
edge of the specimen, so the total number of leaves is greater than 36. The
leaves are all simple and there is no indication of dividing as occurs in Baiera
and allied genera. The tapering of the leaf is very gradual — from 3 4mm.
wide at base to 1 mm. wide at a distance of 12 cm. from the base.

The sporangia vary little in shape, but they may b(^ <livided into two
distinct groups — one in which the individual megaspores are clearly visible,
the other in wLich little or no structiire can be obser\'ed (see Plate I., fig. 2|.
There is some indication in a few of the latter type that the sporangia contain
megaspores, outlines of which can be observed. It may be, however, that
the majority of these sporangia wdiich show no structure now were the micro-
sporangia and that no details of the microspores are preserved in the specimens
available.

The megaspores (Plate I., figs 4. 5), with their equatorial ridge and the
tri-radiate marking on the upper half, are of the type characteristic of Isoetes
and, from the association of the sporangia with the specimen in which the
leaves are so similar to those of species of Isoctes, it is reasonable to conclud('
that sporangia and leaves belonged to the one species and to refer them to
the genus Isoetites. There is indeed evidence that the sporangia occupied the
position usual in Isoetes, several specimens, of which one is figured (Plate I.,
fig. 3), showing a sporangium resting on the wide basal portion of a leaf.

Some specimens, such as that figured on Plate I., lig. 2, show a number
of sporangia (megasporangia and ? microsporangia) arranged more or less
parallel to one another. Some of those on the figurecl specimen have some in-
dication that they rest on portions of leaves.

In size and shape these fossil sporangia are well within the range of those
of recent species of Isoetes^ as also are the megaspores. The range of size of
megaspores in recent species is from 250 to 900 p. Such species as Isoetes
Engelmanni and I. Boryana w'ould bear comparison with our fossils in this
respect (see Pfeiffer, 1922).

There are few recorded fossils that can with certainty be referred to
Isoetites, so that comparison of the Western Australian specimens witli known
species is restricted. The leaves are very different from those of 1. serratus
and I. horridus from Cretaceous rocks in Wyoming, U.S.A. (Brown, 1939, p.
268) and no details of the megaspores are known in these two species.

The fossils described as Isoetites Choffati Saporta from the Lower Cret-
aceous of Portugal are not comparable with our specimens, since they consist
of small tuberous bodies compared with the stem of Isoetes, and imj^ressions
identified as the basal portions of sporophylls bearing sporangia (fide Seward,
1910, p. 67 — a copy of Saporta’s original description does not appear to be
available in Australia).

Fossil Plants from Gingin, W.A.

203

Type speciniPMs iu collection of Geology Department, University of West-
ei*n Australia : Nos. 16687 (leaves) and 16683 (sporangia). Counterparts of
portions of the types are in the collection of the Australian Museum (Nos.
F 39818, F 39816).

FILICALES.

Cladophlebis australis (Morris).

There are one or two specimens referable to Cladophlebis australis. They
are not well preserved and apparently the species is not at all abundant.

Thinnfeidia talbragarensis Walkom.

(Plate II., fig. 8.)

Several specimens agree well with Thinnfeidia talbragarensis from Jurassic
rocks at Talbragar, N.S.W. (Walkom, 1921, p. 9).

The frond is bipinnate and of the type common in Australian Thinn-
feldias but there is no indication that the rachis divides dichotomously as it
does in most Australian spe(*ies of Thinnfeidia.

Sphenopterid fragments.

Several sphenopterid fragments, which show no detail, may belong to a
species such as Coniopteris hyynenophylloides which is common in Jurassic
floras, and is known in association with Taeniojderis spahdata in the Aus-
tralian Jurassic.

Taeniopteris spatulata McClelland.

(Plate II., fig. 9.)

There is a considerable variety of leaves in the collection referable to
Taeniopteris, examination of which only emphasizes the difficulty of accurate
specific definition of numerous sterile Taeniopteris fronds (see Seward, 1904,
p. 169).

These leaves are elongate lanceolate, somewhat strap-shaped, more than
7 cm. long, usually up to 1*6 cm. wide, occasionally somewhat wider (2*4
cm.), with a prominent finely-striated midrib which has a width of 1*5 to
2 mm. The secondary veins are at right angles, or almost so, to the midrib ;
many of them are simple, but many divide, usually only once, at varying
distances from the midrib ; on the average tliere are about 16 veins per cm.
of lamina, but the number \ aries considerably, and in some of the narrower
leaves there may be as many as 25 or 30 veins per cm.

It is not easy to separate these fronds from T. S 2 )atulata McClelland,
and T. spahdata- var. major (Seward). The larger specimens certainly ap-
proach T, Garruthersi Tonison-Woods, a species which has never been very
satisfactorily described, since only incomplete specimens have been available.
The midrib in the Western Australian specimens appears to be much more
prominent than that of T. Garruthersi.

T. spatulata occurs abundantly in the Jurassic rocks of Eastern Aus-
tralia, and occasionally in the Cretaceous. T. Carriithersi, on the other hand,
appears to be a somewliat older form, occurring in the Triassic (Ipswich Series)
of Eastern Australia, and in the 8tormberg Beds of South Africa.

204

A. B. Walkow, l^.Sc.

CYCADALES.

Ptilophyilum pecten (i^iiillips).

(IMatt* II.. %. 0.)

Spccim(‘iis n‘f(‘]T(*(l to J^tilojjhyllum pccien are (juite .‘Similar to tlu‘ many
iimired examples of' tliis spi'cies which is widely distributed in rocks of flurassir
a^e. Tlu'v are i>reser\-e<l as im]>ressions on a fine-grained, h'rruginous mica-
ct'ous sandstotie, and are not likely to furnish any details of cuticnlar structuiv.
It therefore sedans only possibk' to reft'r them to P. pecten (s('C S(‘war<l, 1917, p.
524). In Kastern Australia tliis s})ecies occurs in the Jurassic rocks at Stewart's
Creek. Stanwell. Queensland, and in tlu‘ Cretaceous rocks of the Maryl.)orough
Series and the Ibirrum Series.

CONIFERALES.

Elatocladus plana (Feistinaritel).

(Plate II., fig. 7.)

Several specimens may be referretl to Elatocladus plana Feistmantel. a
common Jurassic type, of which Seward (1919, p. 431) has figured two speci-
mens. The species has also been figuretl from Jm-assic and Cretaceous rocks
in Queensland (Walkom, 1917, PI. 9, fig. 4 ; 1919. PI. 2. figs. 4, 5),

The Western Australian specimens ha\-e linear leaves about 1-5 cm.
long, with a distinct midrib.

LIST OF REFERENCES.

Brown, R. W., 1939 : Some American Fossil Plants belonging to the Isoetales. J. Wash-

ington

Acad. ScL. 29 (6), 261-269.

Pfeiffer, Norma E., 1922 : Monograph of the Isoetaceae. Ann. Missouri

Bo(. Gard.,

X (2).

79-232.

Seward, A.

C.—

1904 ;

On a Collection of Jurassic Plants from \'ictoria. liec. Geol.

Sure. Vicf..

i (3).

p. 155.

1910 :

Fossil Plants, ^'ol. 2. Caml)ridge Univ. Press.

1917 :

Fossil Plants. Vol. 3. Cambridge Univ. Press.

1919 :

Fossil Plants. \o\. 4. Cambridge Univ. Press.

Walkom, A. B. —

1917 :

Jlesozoic Floras of Queensland. Part i. (concluded). Qd.

Geol. Sure.

Paid.

259.

1919 ; Mesozoic Floras of Queensland. Parts iii and iv. Qd. (Jeol. Sure. Pub!. 203.
1921 : Mesozoic Floras of New South Wales. Part i. Mem. GeoL Syn\ A . N. II a/eN.
Palaeontology No. 12.

KXPLANATION OF PhATKS.

Plato 1.

Isoetites elegans, n. sp.

Fig. 1. Isoetites elegans. n. sp. Specimen No. 16687. xj.

Fig. 2. Group of sporangia. Specimen No. 16683. X3.

Fig. 3. Sporangium in position on basal portion of a leaf. Specimen No. 16682. X i.
Fig. 4. Single sporangium. Specimen No. 16711. X4.

Fig. .5. Poi-tion of figure 4 enlarged to show triradiate ridges on megaspoves. x 14.

IM,A'r K

200

A. B. Walkom, D.Sc.

Plate n.

Pig. 0. Ftilophyllum pecteii I^Phillips). Specinieii Xo. 16684. Xl.

Fig. 7. Elatocladus pl-ana (Feistmaiitel). Specimen No. 16703. Xl.
Fig. 8. Thimifeldia talbragaremis Walkom. Specimen No. 16690. xf.
Fig. 9. Taehiopteris ^patuhfa McClelland. Specimen No. 16685. Xl.

Fossil Plants from Gingin, W.A.

207

PLATE II.

Marine Copepods from Western Australia, Series II.

209

11.— MARINE COPEPODS FROM WESTERN AUSTRALIA

SERIES II.

TWO PELAGIC COPEPODS FROM COCKBURN SOUND.

By W. S. Fatrbridoe, B.Sc.

Road 1 4th April, 1942 ; published 27 March, 1944.

The species described were found in some plankton hauls made a few
miles south of Fremantle in Cockburn Sound, about midway between Garden
Island and the mainland. The maximum depth of water in the area is 20
metres.

Order CALANOIDA.

Family CENTROPAGIDAE Sai-s, 1902.

Genus CENTROPAGES Kroyer, 1849.

A large number of ne\^' species ha\ e been described since tViis genus M'as
summarised by Giesbrecht and Schmeil (1898). Unfortunately, the author
has been unable to study' the descriptions of C. longicornis (Mori, 1932) and
C. jjunticus (Karawiew', 1895).

Centropages australiensis sp. nov.

Occurrence. — Regularly found, sometimes forming a considerable pro-
portion of the catch ; males and females in about equal numbers.

Female. — Length, 1*4-1 *6 mm.

The mouth parts agree with those figured for C. typicus (Kroyer, 1848)
by Giesbrecht (1892, Fauna und Flora des Golfes von Neapel) and Sars (1900,
An Account of the Crustacea of Norway'),

The fifth leg differs from C. typicus in the endopod, the second segment
being as long or longer than the terminal and distinctly narrowed at the base,
and the basal segment broadened distally. The second abdominal segment
is very long compared with the genital and anal segments ; the proportions
of the abdominal segments are 23 : 34 : 10 j 18. The genital segment bears
only' two spines, one dorso-lateral and one ventro-lateral, both on the left
side, the latter being very small ; the second segment bears a hooked knob
on its right side. The fifth thoracic segment is only slightly asymmetrical,
the left side being the larger.

Male. — Length, 1*4-1 *5 mm.

The first antenna has a distinct .spine on segments 15 and 10 (Sars does
not figure a spine on segment 15 for G. tyjjicu.^, and Giesbrecht indicates a
minute one) ; thei’e is an incomplete division between segments 22 and 23
(as indicated by Giesbrecht for C. typicus, but vSars show's a complete division).
The other mouth parts agree with those for C. typicus.

The swimming legs appear to be ty'pical : Giesbrecht (1898) says of

C. typicus that the right exopods of legs 3 and 4 bear a “ vergrosserten Aussen-
randdorn ” on the second segment ; this is noticeable in the present species.
The fifth leg differs markcdl}' bum C. typicus in that the basal portion of the
chela is much narrower, resembling the type found in C. hamatus (Lilljeborg,
1853) ; and the proximal segment of the chela extends beyond the “ thumb
and is but slightly hooked at the tip. The abdomen agrees with 0. typicus.
The spines of the last tlioracic segment are asymmetrical, the right being
much smaller than the left.

k 17/43.

210

VV. S. Faikbridge.

llie specimens taken in Coclvbiim Sound appear to be identical with
those found oft Ne^^' South Wales and figured by Dakin and Colefax (1940)
as a variety of C. kroyeri (Giesbrecht, 1892) : the only difl'erence between

these specimens and the rather few figures given for tlie Sydney variety, being
tlie slightly larger basal portion of the cliola of the New South Wales speci-
mens. While the male fifth leg of this species resembles C. kroyeri in some
ways more nearly than C. typicns, the female abdomen and last thoracic
segment are so entirely different from C. kroyeri, that it would seem best to
regard it on these features as more nearly related to C, tyqdcus. Tt resembles
C. typicus rather than C> kroyeri in a mimber of other points : the spines on
the proximal segments of the first antenna ; the outer spine of the second
exopod segments of the third legs ; and the length of the inner spine on the
second segment of the female fifth leg relative to that segment.

On comparison with some drawings made by Dr. A. G. Nicholls, tliis
species appears to be identical also with a Centropagid found by him in 1939
at Crawley Bay in the Swan River. Crawley Bay is well up in the Swan
Inlet, and the salinity would be very low in winter : tliis form is therefore

either originally or secondarily estuarine. Dakin and Colefax give no details
as to where their specimens were found, b\it all or most of their collecting
was done in the open sea. Since therefore, this variety is constant from
New South Wales to Fremantle, in the open sea and under estuarine con-
ditions, it seems best to give it specific rank.

It may be most easily I’ecognised by the female abdomen and the form
of the chela of the male fifth leg.

Order HARPACTICOTDA.

Family TISBIDAE Sars. 1904.

Genus MACHAIROPUS Brady, 1883.

Lang (1936) has revised this genus. Since tlien two new species have
been described ; M. antarcticus (Lang, 1936b) and M. intermedins (Nicholls,
1941).

Machairopus cockburni sp. nov.

Occurrence. — 8 females (3 ovigerous) and 10 immature specimens, in a
horizontal haul at 10 m. depth ; 22-7-41.

Female . — Length, 0 • 94- 1-15 mm.

First antennae 9-segmented, tapering gradually, the proportional lengths
of the segments being 1 8 ; 24 ; 32 : IS : 4 : 4 : 3 : 10 ; 14. Exopod of the
second antennae 4-segmented ; the endopod bearing 9 terminal setae, 4 of
w'hicli are geniculate. Second basal segment of the mandible bears a single,
long, thick, plumose seta ; endopod and exopod typical. Exopod of maxilla
bears 2 long plumose setae ; the endopod with 3 stiff serrate setae ; inner
lobe I with 8 setae, and inner lobes II and III each with 2 setae. First maxilliped
with an inner marginal lobe bearing 2 stout and 1 fine plumose setae, and a
slender distal lobe bearing 2 setae. Second maxilliped with 1 stout claw^
apically and 3 finer setae.

Proportional lengths of the segments of the endopod of the first leg
10 : 6 : 3, each segment with a plumose inner marginal seta ; outer marginal
spine of the second segment of the exopod in the middle of the margin, and
the inner marginal bulge pronounced. Seta formula : —

Endopod. Exopod.

p2 1. 2. 230 1. 1. 223

p3 1. 2. 330 1. 1. 323

p4 1. 2. 230 1. 1. 323

Marine Copepods from Western Australia, Series II.

211

Distal segment of the fiftli leg with almost parallel margins, ami more
than twice as long as wide ; it bears 3 terminal setae, the middle one longer
tlian the other 2 ; 2 external lateral setae, each shorter than the terminals,

the proximal being within the distal half of the segment. Proijortional lengths
of the abdominal segments and caudal rami 73 : 37 : 28 : 1 1 : 1 7. Genital
segment deeply clett ; caudal rami divergent, sliizhtly wider at the base than
long : the outer marginal seta barely within the distal half ; 4 terminal and
1 small dorsal setae.

M ale . — Un known .

This species most nearly resembles M. hippohjtes (Kroyer, 1863),. a
northern cold-water species (Greenland and Norway), and M. australis (Scott,
1912), a southern cold-water species (South Orkneys and South Georgia).
It differs from M. hippoltifes in (a) the relative lengths of the segments of
the first antennae, (b) the setae of the second basal segment of the mandible,
(c) the setae of the inner lobe of the first maxilliped, (d) the relative lengths
of the 2 proximal segments of the emlopod of the first legs, (e) the setation of
the swimming legs and, (f) the shape and setation of the fifth legs. In addition
to the points (a), (b), and (e) above, it difiers from M. australis mostly in
the presence of a distal lobe on the first maxilliped, the shay.ie of the fifth
legs, and the length of the caudal rami to the anal segment (in M. austrahs
“ about as long as the last abdominal segment,” Scott).

It may be recognised by tlie almost parallel margins of the fifth legs,
the proportions of the endopod segments of the first legs, and the single largo
seta on the second basal joint of the mandible.

REFERENCES.

Dakin, W. J., and Colefax, A. N. Plankton of the Coastal Waters off New South Wales,
Pt. I. 1940.

Karawiew, W. “ Contributions to the Copepod fauna of the Black Sea ” (title translated
from Russian). Zapinski Kiev Obskch., XIV. 1895.

Lang, K. “ Uber die Gattungen Psamathe Philippi, 1840 und Machairo()us Brady, 1883,
etc.” Zool. Anz. Bd. 114 Heft 1/2. 1936a.

Lang, K. Swedish Antarc. Exped., 1901-1903, iii, 3. 1936b.

Mori, T. “ New Copepods from the southern waters of Jajjan.” Dobuis. Zusshi Tokyo, 44.
1932.

Nicholls, A. G. “ Littoral Copepoda from South Australia (1) Harpacticoida.” Hec, of
the S. Aust. Mus. Vol. VI., No. 4. 1941.

Scott T “ On the Entomostraca of the Scottish National Antarctic Expedition.” Tran-'-.
Hoy. So'. E(Un., XLVIIL (3). 1912.

212

W. S. Fairbridge.

PLATE I.

Fig. 1 — Gentropages australieyisis.
Fig. 2 . — Gentropages australiensis.
Fig. 3 . — Gentropages australiensis.
Fig. 4 , — Gentropages australiensis.
Fig, 5 .' — Gentropages australie.nsis.
Fig. 6 . — Gentropages australiensis.

? Fifth leg.

? Abdomen (ventral).
cJ' Abdomen (dorsal).

C? Fiftli legs.

$ Abdomen (seen from left side),
? Abdomen (dorsal).

Marine CorKPODS from Western Australia, Series IF.

213

PLATK I.

214

W. S. Fairbridge.

PLATE II.

Fig.

7.-

—Centropages australiensis.

Fig.

8.-

—Centropages australiensis.

Fig.

9.-

—Centropages australiensis.

Fig.

10.-

—M a ch air opus cockhurni .

Fig.

11.-

—Machairopus cockhurni.

Fig.

12.-

— Machairopus cockhurni.

C? Right antenna I.

C? Fourth legs.

C? Third legs (first and second
Exopod Segments).

$ Fifth leg.

5 Abdomen (ventral).

5 Fourth leg.

Marine Coperods prom Western Australia, Series II.

215

PLATE IT.

216

W. S. Fairbridge.

PLATE

Fig-

Fig.

Fig.

Fig.

Fig-

Fig.

13. — Machairopus

1 4 . — Mach air opus

1 5 . — M ach air opus

1 6. — Machairopus

17. — Machairopus

1 8. — Machairopus

cockburni.

cockburni.

cockburni.

cockburni.

cockburni.

cockburni.

$

?

?

?

?

?

III.

Antenna I.
Antenna II.
Mandible.
Maxilla.
Maxilliped II.
Maxilliped I.

Marine Copepods from Western Australia, Series II.

217

PLATE III.

A Consideration of the Insect Population Associated
wiTPi Cow Dung at Crawley, W.A.

219

12.— A CONSIDERATION OF THE INSECT
POPULATION ASSOCIATED WITH COW
DUNG AT CRAWLEY, W.A.

BY G. J. Snowball, B.Sc. (Hons.)*

Read 9th June, 1942 ; Published 28th March, 1944.

1. INTRODUCTION.

The research w^ork described in this paper is an attempt to indicate the
main insects which form the population of cow dung at Crawley and to assign
them to their proper place in the ecology of that substance. Most of the
work was performed at the Biology Department, Crawley, during the periods
February to November, 1940, and February, 1941, to February, 1942. Two
cows supplied most of the dung utilised, but supporting observations at May-
lands and Spearwood showed that this restricted amount of material indicated
correctly the qualitative nature of the population, a conclusion borne out by
the study of insects in tlie dung dropped by a herd of fifteen cow's grazing
in the grounds of the University. The writer also had tlie opportunity of
examining cow dung from Armadale, and in August, 1941, was able to pay
a visit to Katanning to make a rapid comparative sur\-ey of the cow dung
pojoulation.

Katanning (latitude 33° 38^ S., longitude 117° 35^ F.) has a climate
of the continental type, characterised by a considerable \-ariation between
maximum and minimum temperatures. Perth (latitude 31° 52' W, longitude
110° 3' F.) has a Mediterranean climate with less difference between the ex-
tremes of temperature. The rainfall at Katanning, although annually amount-
ing to only about half that at Pei'th, is much more evenlv distributed through-
out the year, rainfall at Perth being essentially of the winter type.

Table 1 .

The mean daily maximum and minimum temperatures, the mean daily
relative humidity, and the mean annual rainfall at Katanning and Perth,
Western Australia, for 34 years prior to 1932 (based on figures from the
Weather Bureau, Perth) :■ —

PERTH.

katanning

Maximum ° C.

22-89

21-94

Minimum ° C.

12-89

9-11

Humidity

03

70

Rainfall (inches)

34-7

18-7

2. MFTHODS,

No accurate t[uantitative work was attempted in tins general approach
to the problem. The insects found in various cow dungs were submitted to
experts for identitication where possible and obsc'r\ations made on their
habits. Portions of cow dung containing immature stages were brought
into tlie laboratory for study of jihases of the life histories of the insects con-
cerned.

3. SUCCFSSION.

A large numlier of insect species are associated with faecal matter,
cs])ecially with that of mammals, which is usually of sufficient ({uantity to
form an ecological habitat of some duration.

*In 1941 tlie writer was the recipient of a Ilackett Scholarship, and in 194 ^ of a
Hackett Studentshij>, for which his grateful thanks are due to the University of W.a"

220

G. J. Snowball.

In an imstable environment of a restricted nature, of which cow dung
is an example, two main factors operate to influence the type of population
present at any particular time. Firstly, the external climatic conditions
change the nature of the environment, thus making it less suitable for the
species which occupies it first. Secondly, the animals themselves, principally
by interspecific competition, to a lesser extent by altering the physical and
chemical nature of their environment, tend to bring about an alteration in
the type of population. Ultimately a stage is reached at which a primary
species is displaced by one more adapted to the changed conditions. The
process continues until the environment is destroyed. This termination of
the succession by destruction of the environment, which is characteristic of
the animal ecology, serves to balance, in the broad cycle of energy of nature,
the synthetic tendencies of plant successions in which a stable association is
finally attained (Chapman, 1931).

The phenomenon of succession has received most attention from plant
ecologists. The most clear cut examples of succession in animal ecology are
found in certain restricted environments. The work of Graham (1925) on
the succession of animals in a felled tree trunk may be cited here. He con-
cluded that there was a definite succession of organisms as the chemical and
physical characters of the wood changed during the process of decay and dis-
integration. Fuller (1934) found this statement to be equally true of the
population of a carcass.

In the case of cow dung, the succession is not clear cut and only detailed
study would reveal the causes of some apparent anomalies. The nature of
the succession will be discussed after the insects concerned have been dealt
with.

4. THE CHANGES IN COW DUNG AFTER DROPPING.

The initial nature of a cow dung varies considerably with the diet of the
animal. The dung of a grass-fed cow differs from that of one fed on dried
feed in its green colour, soft texture, and the subsequent course of its dis-
integration. The process of decay can be arbitrarily divided into five stages,
of which the first three differ markedly from the last two. A characteristic
insect population occupies the first three stages and a different one succeeds it
in the last two. Certain distinctions can in many cases be dravn between the
populations of Stages 1, 2, and 3, but equally often they are almost identical.

Following is the typical course of changes in dvmg from a grass-fed cow : — ■

Stage.

When dropped, the dung is green or more rarely tan in colour, semiliquid,
and with a noticeable odour. Cow dung is characteristically a cold ” dvmg
and liigh temperatures due to fermentation are unusual, especially in isolated
cakes, although the temperature of the dung during the day may be high
owing to exposure to the sun (Davidson, 1937). The slight degree of fer-
mentation undoubtedly influences the insect population since certain insects,
e.g., tlie house fly, are specifically attracted to fermentable materials either
by high temperatures or products of such decomposition.

The first obvious change in the cow dung is the formation of a thin
blackish skin over the exposed surface of the soft mass. By evaporation
of moisture, this skin ultimately becomes a thick, firm crust pierced by many
cracks which expose the softer interior. The superficial skin is the result of
an oxidation process dependent on free access of atmospheric oxygen, since
it does not form on artificially covered cakes or those which for any reason
are kept exceedingly moist.

A COXSIDEKATIOX OF THE INSECT POPULATION ASSOCIATED

WITH Cow Dung at Crawley^ W.A.

221

2nd Stage.

The colour changes from tan or green to a light brown shade. The whole
cake becomes firmer as a result of evaporation of moisture. The odour is
still noticeable even when the surface remains unbroken.

3rd Stage.

The dung becomes blacker and firmer in texture. The fibres of the*
undigested food can be distinguished, because evaporation of moisture has
concentrated the suspension of which they form the discontinuous phase.
The odour is usually not noticeable unless the surface of the dung is disturbed.

4:th Stage.

Evaporation of moisture has proceeded to the extent that undigested
food fibres interlock to form a matrix firm enough for the whole cake to be
held by the edge without breaking. The internal colour is a dull rusty brown.
The surface layers of the dung are completely dry and of a bleached brown
or even white colour due to the precipitation of salts.

^th Stage.

The entire dung becomes of the same nature as the upper portions of a
fourth stage dung. It is firm, bleached brown, in some cases almost white,,
dry, hard, and noticeably light for its volume. In the later part of this stage
no insects except casual shelterers are present.

A number of variations from the above cycle are possible. All parts
of a dung cake do not necessarily pass through the same stage at the same
time. For instance, dungs which have the upper layers in the fourth, and
lower portions in the second or third stage, are common. Further, the changes'
in colour, texture, and moisture do not always proceed at the same rate.-
Dungs sometimes preserve the initial green colour although in other respects-
they could be classed as in the fourth stage. Such peculiarities are due to-
some unusual feature in the history of the cake, such as freedom from invasion,
by insects or conditions which artificially preserve the moisture content.

An important departure from the cycle outlined above is the omission,
of either the second or the third stage. The colour of a dung may pass straight
from green to black, i.e., direct from first to third, or from light brown to dark
brown, i.e., from second to fourth. In such cases, the population of the-
omitted stage is found in the early part of the third or late jmrt of the second
according to whether the second or third has been rt'duced.

In the field, large numbers of dungs disintegrate before they reach the
fifth stage, an occurrence very frec|uent during tlie height of the summer..

The time taken to pass through the \-arious stages of decomposition*
varies greatly with climatic conditions, so that, exceiit witliin wide limits,,
the absolute age of the dung has little significance. High temperatures'
speed up the process of disintegration by accelerating the cliemical and physi-
cal processes within the dung and by stimulating the contained insects. Rain
has the reverse effect, and the stages of a dung ex 2 iosed to rain may be indefin-
itely prolonged.

-222

G. J. Snowball.,

The following table indicates a few variations in absolute time of stage
<iuration for various conditions of rainfall and temperature : — •

Table 2.

Duration in days of each separate stage numbered.

Date of
Dropping.

1.

2

3.

4.

5.

Rain during
period of
disintegration.

Temperature
during period of
disintegration.

.20-3-41

18

1 3

7

*

Nil

20-6° C.

J2-4-41

2

t

8

15

*

2-84"

18*0'^ C.

7-4-41

1

4-

+

H

10

*

1*50"

17-6° C.

31-5-41

4

8

14

*

. .

7-07"

14-2° C.

4-6-41

8

7

29

40

*

20-00"

13-9'^ C.

12-6-41

9

t23

. . .

41

*

17-14"

13-9° C.

22-6^1

7

18

*

. • •

. « «

5-52"

14-2° C.

23-7-41

8

3

29

*

...

6-86"

13-9° C.

-* Broken up in this stage. f -3 Duration of stage 2 plus 3 because they could

not be separated. J Omitted.

Insects are rarely responsible for tiie comj)lete destruction of a cow dung,
this being accomplished by rain and other mechanical agencies. On the
•other hand, c*akes may remain in the fifth stage for years without noticeable
i^rosion.

The cycle in a dung heap is substantially the same as that in a single
cake, although the duration of the stages is usually greater because the inner
portions are less accessible to disruptive influences.

5. THE INSECTvS CONCERNED.

The following is a list of the Insect Species associated with cow dung at
Crawley, Western Australia : —

CoUembola

r : Hypogastruridae
* Uuidentifled specimens

Derniaptera

F : n.,a!>iduridae

Unidentified sjiecies

Orthoptera

F : Tettisoniidae

Unidentified si)ecimens
F : Gryllotaliddae
Cj liiniracheta s\).

ilemiptera

F : Pentatomidae

Unidentified sj)ecimens
F : (’occidae

Unidentified specimens

Coleoptera

F : Uarahidae

Promecodcrus albanyensis, Cast.
y : Hydropliilidae

* Uei’cyon haemorrlioidalis. Fai).

* Cercyon nicriceps, Marsh.

V Parac'ymus sp.

F ; Sta])hylinidae

Ureophilus erythrocejfiialus, Fab.

* Ury])tobiiim elegans, Blkb.

* Le)itacinus socius, Fauv.
Philonthus sp.

Oxytclus spp.

F ; Silphidae

V Ciioieva sp.

F : Tiichopterygidae

V E))optia sp.

? Philagai’ica sj).

F : Histeridae

Saprimis incisiis, Er.

Saprimi.*< s])p.

Hi.ster walkcri, Lea.

Piatysoma ? nmltistriatum, Lea.

1*' : Nitidulidae

Uarpo])hiliis hemii)teriis, Linn.

? Brachyjjeplus sp.

F : Colydiidae

Pabula dentata. Blkb.

F : Ptinidae

? Ptinus sp.

* Breeding in cow dung.

A COXSIDERATIOX OF THE lX.SECT POPULATiOX ASSOCIATED

'WITH Cow Dhxg at Crawley^ W.A.

22‘3

¥ : Scarabaeidae

* Aphodius lividus. Oliv.
Aphodius insignior, Blkb.
Aphodius granarius, Linn.
Aphodius ambiguus, Boh.
Aphodius sp.

* Proctophanes sculptus, Hope.
Ataenius ? integricollis, Lea.
Onthophagus ferox, Har.

F : Anfrhicidae

Anthicus hesperi, King.

F : Tenebrionidae

Gonocephalum arenarium, Fab.
Adelium scytailicum, Base.

Diptera

F ; Psychodidae
* ‘ Psychoda spp.

F : Sciaridae

* Sciara spp

* ? Zygoneura sp.

F : Scatopsidae

? llhegmoclema sp.

F : Mycetophilidae

? Mycetophila sp.

F : Tipulidae

Eriopterinae

* Unidentified species
F : Stratiomyidae

Xeoexaireta spinigera, Wied.

F : Dolichopodidae

? A.syndetus sp.

F : Phoridae

? Chaetocnemistoptera sp.

F : Sepsidae

* Sepsis plebeia, de Meij.

* Australosepsis fulvescens, Mall.

* Australosepsis fulvescens var.

atratula, Mall.

F : Borboridae

* I,eptocera spp.

F : Drosophilidae

* ? Cladochaeta s]).

F : Otitidae

* Chrysomyza aenea, Fab.
Pogonortalis barbifera, Macq.

F : Calliphoridae
Calliphorinae

Calliphora (Neopolienia) australis,
Boisd.

Calliphora (Proeckon) nociva.
Hardy.

Lucilia cuprina, Wied.
Sarcophaglnae

* Sarcopiiaga (Para.sarcophaga>

depressa, Desvoidy

F : Miiscidae
Muscinae

* Musca vetustissima, Walker.

* Musca domestica, Linn.
Anthomyiinae

* Hylemyia deceptiva, Mall, or

near

* Hylemyia sp.

Fanniinae

* V Fannia sp.

Phaoniinae

* Helina hypopleiiralis, Mall, or

near

* Helina coerulescens, Stein.

* Helina regina, Mai. or near

* Rhynchomydaea (Hardyia)

carinata, Stein or near

* Muscina stabulans, Meig.

Lepidoptera

F : Oecoi)horidae

* Unidentified spec’es

Hymenoptera
F : Alvsiidae

1 unidentified species

F : Figitidae

1 unidentified species

F3 : Formicidae

Unidentified species

6. RELATIONS OF INSECTS TO THEIR ENVIRONMENT AND TO>

EACH OTHER.

These may be outlined as follows ; —

). Those which do not live in the cake itself but use it as food,
blowflies. Coprine beetles.

2. Predators on duns: insects with no closer reiationsh’p to the dnng-

Tho.se iisiially remain on the outside of the cake, cxy., Dolichopo-
(lids, Creophiius erythrocephalus^ and ants.

3. Those which live in or on the dung, but subsist on the fungal growths

which it supports. Probably the minute Collembola frequent-
ing dung have this habit, although their small size renders
the elucidation of this point very difficult.

4. Insects, the larval stages of which live in and feed upon dung

while the adults have other feeding habits, e.g., some Sarco-
phagids and probably Neoexaireta spinigera.

5. Insects which are predatory themselves or have predatory larvae-

and of which the whole life history is passed within the dung,
e.g., Oxgtelus, Leptanmis, Cryptobium, Saprinus, and Platysoma.

Reared from cow dung.

224

G. J. Snowball.

6. Insects which use the dung as a shelter, e.g., Pentatomida, Dermap-

tera, and Tenebrionids, the association of which with the rest
of the cow dung population is only of the most casual nature.
Ants are in this class but they are also predatory.

7. The true dung insects, which are coprophagous and pass through

their entire life history in the dung, e.g., the Aphodii. Owing
to a doubt concerning the food of the larvae of the Cercyons,
it is not known whether they can be classed here.

8. Parasites of dung-living larvae, e.q., Aiysiids and Figitids.

7. NOTES ON THE INSECTS ASSOCIATED WITH COW DUNG AT

CRAWLEY.

Order l—COLLEMBOLA.

Myriads of minute blue insects belonging to the Hypogastruridae occur
in and on dung during the winter, June to August. Large numbers occur
in any stage of the dung providing it has remained fairly moist.

The wdiole life history can apparently be passed through in cow dung.
Eggs are fairly common in the dung, most noticeably so in June. Some
brought into the laboratory in June, 1941, hatched in twelve days, but the
insects were not raised to maturity. At times, parts of first and second stage
-dungs in the field are almost wdiite with the exuviae of these tiny insects.

Order 2— DERMAPTERA.

Unidentified specimens of Labiduroid Dermaptera are occasionally
-shelterers under cow dungs in the field. The association with the dung appears
.to be nothing more than casual.

Order 3— ORTHOPTERA.

Tettigoniid nymphs are sometimes found sheltering under old cow dung,
•while a Cylindracheta species found at Crawdey is apparently capable of a
truly coprophagous habit, since two specimens lived for some weeks on a diet
.of pure COW’ dung.

Order 4— HEMIPTERA.

At Katanning in August, 1941, a number of Pentatomids w’ere collected
in crevices of very old fifth stage dungs wLich they used as a shelter only.
The same is true of Coccid nymphs found under old dungs at Spearw’ood,
Western Australia, in April, 1941.

Order 5— COLEOPTERA.

F. 1. CARABIDAE.

Promecoderus albanyensis, Cast.

Specimens of this wingless beetle occur under cow’ dung at Crawley. It
is presumably predaceous but its timidity has so far prevented observation
of its habits.

The larvae of unidentified Carabs are also common under, and more
rarely in, dung cakes. One was kept alive during August and September,
1941, on a diet of Sepsid i . 1 Sciarid larvae, but died before pupating.

F. 2. HYDROPHILIDAE.

The genus Cercyon contains both aquatic and terrestrial species, many
of the latter being inhabitants of dung, Stephens (1839) having described

A Consideration op the Insect Population Associated
WITH Cow Dung at Crawley^ W.A.

225

42 species with this habit. Sphaeridium, a coprophagous genus wliich I iiave
not seen recorded from Australia, is associated with Cercyon in both England
and Prance, especially in cow dung (Lacordaire, 1854), and in Ceylon and
Japan aquatic species of both occur (Sharp, 1884, 1890).

Cercyon haemorrhoidalis. Fab.

This introduced species is represented at both Crawley and Katanning
by two easily separated varieties. One is most numerous between November
and June (PI. 1, Fig. 1), the other between Juno and November, both being
present all the year round. The species occurs also in carrion, decaying
grass, and horse manure.

With the Aphodiines, Staphylinids, and Histerids, they arrive at the
dung soon after dropping, tlie odour being apparently the attracting stimulus.
At first they congregate on the under surface or in the natural cracks of the
dung. From these vantage points the work of penetration begins. The
adults are coprophagous, and tunnelled out dung is eaten, since no waste is
found outside infested cakes. The area of penetration is peripheral at first,
then the insects concentrate on the upper surface. Many dungs are found
with an almost unaltered' core, though the superficial layers are completely
penetrated underneath the firm outer crust which is pierced by numerous
holes.

Under favourable conditions of fine days anti high temperatures cakes
may be riddled into a bran-like mass in a single day. Whore heavy rain has
fallen or low temperatures prevail, the process of penetration may bo greatly
slowed down, and may not even occur to any extent at all.

The specimens of Cercyon haonorrhoidalis run into dark places only when
a dung is disturbed, and are otherwis(> tolerant of light.

The beetles leave the dung after a varying period, which is generally
from four to fourteen days. It may be mucli longer and fluctuates con-
siderably from cake to cake under identical external conditions. During
this period, which is one of tro])liic activity, eggs are deposited, since larvae
begin to appear in the third stage dung. I have not been able to isolate tlie
eggs of the species in spite of the larvat> being fairly plentiful. Numbers of
females caged with dung in tlie laboratory died without oviposition, yet
dissection showed their ovaiies contained numerous oocytes.

Exactly what the legless mandibulate larva (PI. l,Fig. 2) feeds on is a
problem. The gut contains a translucent reddish-brown lk|uid full of oil
globules which cannot certainly be recognised as a derivative of cow dung.
Development occupies approximately 10 to 14 days in pure cow dung.

Pupation generally occurs in the dung itself in a small cavity excavated
by the propupa near to the outer surface. Wluu'o the dung lias remained
very moist, pupation may occur in the sand below the dung. No cocoon of
any sort is formed. The jiuj^ae are of the usual exarate coleopterous type
and, like many other white or yi'llow immature stages of insects, is negatively
phototropic.

The pupal period was observi'd to range from three days in April (18° C.)
to II days in .September (18-9° C'.). The life cycle from egg to adult took
less than 29 days at temperatures from 13*9° C. to 10-1° C. in September
anti October, 1941.

220

G. J. Snowball.

There is an indefinite number of pjonerations per year, as larvae and pupae
occiu- in numbers at Crawley for most of the year except for a marked pause
in June and July, when only occasional ones are found. Owing to the com-
paratively short diiration of stages suitable to Cercyon larvae, a second genera-
tion never develo])s in any one dung cake.

Cercyon nigriceps, Marsh.

This small species is associated with C. haemorrhoidalis at Crawley and
Katanning, though always in small numbers. Its life history and habits are
similar to those of its larger relation.

? Paracymus, sp.

A number of small beetles taken in cow dung at Crawley in April, 1941,
probably belong to this genus. Their rarity indicates that they are of little
importance in the dung community.

F. 3. STAPHYLINIDAE.

Creophilus erythrocephalus. Fab.

On warm days at most times of tlie year specimens of this beetle gather
on fresh dung cakes to catch Sepsids and llorborids attracted to it. Usually
they remain on the outside of the cake, but may retreat into cracks if dis-
turbed, or oven hollow out cavities on its lower surface. According to Fuller
(1934) they attack immature stages of sheep blowflies breeding in carrion,
without appearing in sufficient numbers to have much effect on the maggot
population of a carcass. This is also true of the dung cake, as rarely more
than six specimens will be found on the one pad, which means that only a
small percentage of the myriads of flies whicli foregather are destroyed. C,
erythrocephalus rarely remains after tlie decline in number of flies consequent
on the formation of a superficial crust on the cake. Because of this it is not
known whether it will attack dung-inhabiting larvae.

Philonthus, sp.

An unidentified species of tliis genus is occasionally found in small
numbers, usually in second or third stage cow dung. It is more numerous
in decaying grass. Fuller (1934) states that P. Folitus, Linn, eats eggs and
young larval blowflies in carrion. The genus is common in England, where
four species are recorded by Stephens (1939), mostly in dung, and it is repre-
sented in New Zealand (Broun, 1880).

Leptacinus socius, Fauv,

This species, which is presumably predaceous on small Dipterous larvae,
is present in numbers less than the Cercyons throughout the year in cow dung
of all stages, mainly in the first and third. It was not seen at Katanning
in August, 1941. Although Staphylinid eggs and larvae arc common in fourth
stage dung, attempts to obtain adults failed. The larvae of Leptacinus socius
may leave the dung to pupate because in September, 1941, pupae were found
in the sand under an 81 day old dung kept in the laboratory since dropping.
Dung from Armadale collected in March, 1941, contained a pupa (PI, 1, Fig.
3) of this beetle actually in the dung itself.

The pupal period under laboratory conditions was found to be in the
region of 26 days at an average temperature of 13-9° C. in September, 1941.

A Consideration op the Insect Population Associated
WITH Cow Dung at Crawley^ W.A.

227

Cryptobium elegans, BJkb.

This species is associated with L. socius^ but nothing of its life history is
known. The genus is represented by 20 species in England, 19 of which
live in dung and similar material (Stephens, 1839).

Oxytelus, sp.

Several unidentified species are numerous in cow dung, sometimes greatly
exceeding Leptacinus and Cryptobium in numbers. They are among the
first to colonise freshly dropped dung, especially if it has been scattered in
a number of shallow pieces. They also occur in numbers in carrion. In
northern Australia they may play a part in reducing the incidence of the
Buffalo Fly (Lyperosia exigua, de Meij) by competing with its larvae in cow
dung (Mackerass, 1932).

The only immature stage seen was a single exarate piipa in September,
1941, in the sand under dung, three months old, kept in the laboratory since
dropping.

F. 4. SILPHIDAE.

Only two specimens, probably belonging to Choleva sp., were collected
from third stage cow dung at Crawley in March, 1941, and hence the family
is unimportant in the succession. Two species of Choleva occur in carrion
(Fuller, 1934).

F. 5. TRICHOPTERYGIDAE.

At Crawley, species near Epoptia and Philagarica in cow dung in all
stages, especially in the fourth if it is not too dry. Their habits are not kno\vn.
The family is best represented in the tropics, occurring in rubbish, leaves,,
under bark or in rotten wood (Lefroy, 1923).

F. 6 . HISTERIDAE.

Members of this family are present in cow dung at Crawley between
October and May, although they are found in carrion and decaying grass at
other seasons of the year.

SAPRINUS.

Two species of this genus occur in cow dung of which only one, S. incisus
Er., has been identified. Both exhibit a preference for fresh cow dung and
tolerate excess moisture better than do the Hydrophilids.

They never remain in the dung beyond the third stage, which is the one
containing the majority of their prey, larvae of Sepsids, Drosophilids, and
probably Borborids. Small quantities of dung are also eaten. Three larvae
were found in fourth stage dung brought from Armadale to St. George’s Col-
lege in March, 1941. One of those subsequently pupated ; the adult emerging
after three wrecks at an average temperature of 18° C.

The genus Saprinus, like other insects in decaying materials, is widespread.
Species of it occur in England (Stephens, 1839), New' Zealand (Broun, 1880),
and in carrion at Canberra (Fuller, 1934).

PLATYSOMA.

One specimen of P. ? nmltistriatmv , Lea, w'as collected in second stage
cow dung on 4th March, 1941. Stephens (1839) describes three species in
England all under bark.

228

G. J. Snowball.

HIST R.

H. walkeri, Lew,, is the largest of the dung-frequenting beetles at Crawley.
It is most numerous from October to April. It is frequently found in quiescent
state in the moist sand below a cake and is not often found inside. In this it
differs from coenosus, which in Haiti is such an efficient predator on Muscoid
maggots in fresh moist dung that Myers (1938) has recommended its intro-
duction into Australia as a possible control of the Buffalo Fly.

The local species is also predatory on maggots. It probably takes up its
■characteristic position in order to catch maggots which come to pupate on
the lower surface of the cake. Unlike Saprinus, it may occur with fourth stage
dung.

Hister is represented in England and New Zealand.

F. 7. NITIDULIDAE.

One specimen of Carpophilus heniipterus^ Linn., which in California is an
important pest of fresh and dried figs (Simmons, Reed, McGregor, 1931), was
collected from second stage dung in September, 1940. If duo to anything but
chance, its presence indicates a considerable departure from the usual habits
of the species. In April, 1940, two specimens of ? Brachypeplus sp. were also
found in dung of the same stage.

F. 8. COLYDIIDAE.

Three specimens of Pahula dentata BIkb. were obtained from cow dung in
April, 1941, The members of the family are generally found in leaf mould,
decaying w^ood or under bark (Tillyard, 1926).

F. 9. PTINIDAE.

In May, 1941, a single specimen of ? Ptinus sp. was found in a dry, bleached
fifth-stage pad. Its presence, if not merely forti itor.s, adds yet another food
material to the remarkable list given for the family by Esdaile (1927)— farina-
ceous matter, drugs, books, tobacco, spices, pepper, and w^ood.

F. 10. SCARABAEIDAE.

APHODIUS.

Aphodius lividus Oliv.

This species has a w’orld-wdde distribution (Schmidt, 1922). With Ataenius
stercorator it is abundant in Puerto Rico, wfiiere the two eat so much of the
fresh cow dung that the larvae of the Horn Fly (Lyperosia irritans) can scarcely
live. (Observations quoted by Myers, 1938). Myers, liowever, states that
in Haiti these beetles have little effect on the population density of tlie Horn
Fly. A. lividus occurs in large numbers at Uvalde, Texas, U.S.A., wffiere
1,113 specimens have been collected from a single dropping (Lindquist, 1935).
Though not yet finalised, sampling counts at Crawfiey indicate that greater
numbers per cake are not unusual here. Stephens (1839) includes it in his
list of fifty-seven English species of Aphodius, nearly all of which occur in dung.
Boucomont (1929) reports its presence in dung in China. At Crawdey it forms
a prominent part of the population of both horse and cow' dung and is present
at Katanning. At Crawley it appears at and enters the dung shortly after
dropping, leaving again at some time in the third stage. All feeding stages are
coprophagous.

A Consideration of the Insect Population Associated
WITH Cow Dung at Crawley, W.A.

229

The actual information on the life history of A. lividus is slight and much
of it must be conjectured. This is due to the slow growth of the larvae and the
difficulty of handling them imder laboratory conditions. Eggs and larvae of
Scarabs are numerous in cow dung at Crawley, but strangely enough most of
them are those of Proctophanes sculptus Hope, the adults of w’hich are much
fewer in number than those of A. lividits.

Scarab eggs are common in fourth stage dung from February to August.
Eggs have been kept in the laboratory up to 11 days before liatching. This
represents less than the minimum egg period since none were actually col-
lected immediately after laying. Altho\igh Scarab larvae are adapted to the
firm, rusty-brown dung of the fourth stage, they sometimes occur in the moister,
■fresher dung of the third.

The life history period from the egg to the emergence of the adult was
foimd to be as short as 45 days during March (average temperature 20*6° C.)
and April (18*6° C.), 1941. Three larvae which pupated in April emerged in
May after pupal periods of fourteen to sixteen days. It seems that under cer-
tain circumstances the period of development from the egg to the adult can be
considerably shorter than that of A. tasmaniae, which is approximately one
year (Swan. 1934). Lindquist (1935) estimated the life history period of A.
lividus to be 25 to 45 days at Uvalde, U.S.A. under laboratory conditions.

Pupation usually occurs in the sand below the dung., the pupa (PI. 1,
Fig. 4) being unprotected by any cocoon structure.

It is probable that the species breeds in other media besides cow dung,
•since the larvae seen were very few compared with the large number of adults.

Aphodius insignior, Blkb,

Blackburn (1904) erected this species from specimens taken near the Swan
River, and Schmidt (1922) only gives Western Australia as its area of distribu-
tion.

No representatives were found at Katanning in August, 1941. At Crawley
they show a remarkable seasonal distribution. Up till June, 4, 1941, no speci-
mens had been collected since August, 1940. On that date one specimen was
discovered in a second stage cow dung. On 16th June, following rain, myriads
appeared, swarming in fresh dung near tlie Biology Department. They prac-
tically excluded all other beetles and burrow'ed even into the most liquid cakes.
The large numbers were maintained until the end of July when a decline set in.
By 30th August they liad again disappeared.

This appearance in large numbers suggests that A. insignior has a life
history similar to 'that of A. tasmaniae j which has an annual emergence of adults
over a short period of the year. As no immature stages of A. insignior have so
far been found, this must remain conjectural for the present.

The species exhibits a marked preference for fresli, greenish dung on which
it feeds freely. The beetles are exceedingly tolerant of moist conditions, which
gives them a considerable advantage over other insects in colonising cakes.
When inside the dung or on it they habitually associate in pairs. Hanging
on to the prothorax of the female with its forelegs, the male is carried round on
her back and requires considerable persuasion to dismount. This habit is
no doubt a prelude to mating, though actual copulation has not been observed .

A. insignior rarely remains on the dung beyond the second stage.

230

G. J. Snowball.

Aphodius granarius, Linn.

This world-wide species is present botli at Crawley and Katanning in horse
and cow dung. Its small numbers preclude its being of much importance
in the succession.

Aphodius ambiguus, Bohexn.

Also kno^\^l as A.frenchi Blkb, this species is widely distributed in Africa
and Australia (Lea, 1923). Its importance in the dung community v^aries
greatly at Crawley and Katanning. At Crawley, specimens are most numerous
in the period May to August, the total range being from mid-April to October.
Their numbers are always far short of those of A . lividus or G. haemorrlioidalis*
Tliey aj^pear beneath the fresh dung, where they remain until the majority
of the other dung-invading beetles have left. By this time the dung has
usually attained the fourth stage. The beetles then commence to migrate
inside. This well marked succession of the species to the others is a feature of
the winter cycle in dung. In many cases they remain c^uiescent on the lower
surface without entering the dung. While they have never been seen feedings
the gut contents indicate that dung forms the staple food. At Crawley they
also frequent rotting bark, carrion, and decaying grass.

At Katanning in August, 1941, a different state of affairs was seen. In-
stead of being in the minority, they far exceeded all the other dung-frequenting
beetles in number. As at Crawley, they tended to remain on the low^er surface
of cakes of all stages though numbers had penetrated even into fresh dung.

Nothing is known of the life history of A. awbiguus in spite of the presence
of numerous females vith well-developed oocytes in their ovaries. Possibly
some of the Scarab larvae found in the cow dung belong to this species.

Proctophanes sculptus, Hope.

This bulky beetle occurs all the year round at Crawley, becoming most
numerous in the period May to August. Its position in the succession is
difficult to estimate as it frequently occurs associated with A. ambiguus but
also singly in very fresh and more rarely in dry fifth stage dung. Both larvae
and adults are coprophagous.

Owing to the length of the life history and the fierce competition offered
by Sciarid larvae under laboratory conditions, the information on it has been
collected only from a few cases. Seven specimens were reared from the egg
during the period June to November, 1941, and the length of the period from
hatching to emergence of the adult ranged from 97 to 157 days.

The eggs, which are scattered singly in fourth stage dung, hatch about
12 days after laying. The pupal period is approximately three weeks, the
pupae being found in the sand under the dung. The lar\'al period is thus shorter
than that of Aphodius tasmaniae (Swan, 1934), which takes from April to De-
cember. The larvae Proctophanes may remain in a quiescent non-feeding
state for as long as 48 days prior to pupation.

Ataenius ? inTegricollis, Lea.

In August, 1940, three specimens were collected in cow dung at Crawley,,
the species having been described from Queensland (Lea, 1923B). Specimens
of an unidentified species of Ataenius, two at Crawley in December, 1941, and
one at Dog Swamp, Tuart Hill in January, 1942, were taken from cow dung.
According to Blackburn (1904) tlie Australian species of Ataenius are probably
numerous. Considerably more of them came to his notice than all the described

A Consideration of the Insect Population Associated
■WITH Cow DunCi at Crawley^ W.A.

231

Apliodii of Australia at the time of writing: his papei’. In view of this state-
ment and the association of species of Ataenius with Aphodius Uvidus in other
parts of the world (Myers, 1938 ; Lindquist, 1935), the i)aucity of Ataenii
at Crawley is surprising. None were found at Katanning in August, 1941.

Sub. Family COPRINAE.

Although they play no part in the succession in cow dung, Chprine dung
beetles are of inten^st because of their habits, whicli have made them the
subject of study from the earliest times. The ancient Egyptians regarded the
manipulations of circular dung pellets by ScarahaeMS sacer as a symbol of the
revolution of the planets, and its periodic appearance and disap])carance as
a sign of eternal life. Tlie Coprinao utilise dung as food for themselves and
their larvae without actually living in it. This interesting grouj) is not as
well represented in Australia as elsewhere and the species in general are smaller
(Lea, 1923A). The chief Western Australian genus is Onthophagus, which
contains 10 English species found in dung (Stephens, 1839), while Lea (1923A)
lists 102 Australian, of which two, O. nitidor Blkb. and O. australis Guer.,
fre(]uent carrion at Canberra (Fuller, 1 934). One specimen only has been found
at Crawley, but 0. ferox has been taken at Belmont and Clanunont, while in
the country it occurs at Katanning and at Capel.

Onthophagus ferox digs vertical shafts in close proximity to or under cakes
of dung. At about a depth of four inches the direction changes to the hori-
zontal, the resulting passage being about three inches long. In the blind end
of this the store of dung is placed. Fresh pig, sheep, cow or horse dung is
used and stored in the form of scraps, not moulded into pellets. The excavator
of the tunnel is usually found at the bottom of the vertical shaft during the
day, the digging apparently being performed at night. There was no indication
that the beetles work in pairs as is the case with Copris lunaris (Wheeler, 1922).
The presence of Onthophagus is indicated by piles of turned-up earth round
the edges of dung cakes.

F. 11. ANTHICIDAE.

In March, 1941, two specimens oi AnthicMS hesperi. King, were collected
from cow dung at Crawley. This maintains the relationship between carrion
and dung insects, since A . hoejeri, Kerg. is a carrion insect at Canbt^rra (Fuller,
1934).

F. 12. TENEBRIONIDAE.

Gonocephahim arenarium, Fab. and Adelium scytallicum. Base., are ])resent
at Crawley as shelterers under old, drk'd liftli stag(i cow dungs.

Order ti^DIPTERA.

F. I. PSYCHODIDAE.

These tiny moth-like flies are commonly associated with decaying veg(‘tablo
matter, dung or water. Some species of Psychoda breed in drain i)ipes, the
larvae being able to survive hot water and soap (Curran, 1934). The family
is widespread. Pertcoma has been bi'c'd from horse and cow dung in Denmark
(Thomsen & Hammer, 1930), Psychoda 'minuta from cow dung at Washington
(Howard, 1912). Two species of Psychoda frefjuent carrion at Canlx'rra
(Fuller, 1934). The economic imjDortance in Australia is negligible apart from
occasional nuisance caused by large numbers getting into exposed food material.

232

G. J. Snowball.

At Crawley there appeared to be two species of Psychoda, a common
grey one and a rarer wliite. From May to October the larvae, pupae, and
adults of grey species occur on and in soft, moist, first to third stage cakes,
especially those in the shade. Botli larvae (PL 1. Fig, 5) and pupae have
respiratory siphons which are apparently responsible for the high moistiire
toleration.

At the time of pupation the larvae come to the surface. The cracks of
a cake may bo packed tight vith the naked pupae. The total time of life
history from egg to emergence of adult was 6|- days in September, 1941 (average
temperature, 13*9° C.).

Tlie Psychodidae breed in fresher dung than the Sciaridae and con-
sequently precede them in the succession. In the same stage dung as Psycho-
did lar^-ae and pupdC, Sepsid and Drosophilid larva occur, followed later by
Cercyon larvae.

F. 2. SCIARIDAE.

#

Though small flies belonging to Sciara sp. infest dung in the laboratory,
they are not numerous in the field. The genus Sciara is world-wide, having
been recorded from Korth and South America, Africa, Southern Asia, and
Kew Zealand (Skuse, 1889). Tonnoir (1929) estimated the number of de-
scribed species of Sciara and those in collections in Australia, at 63.

They can breed in a wide range of organic materials. Fuller (1934).
records them from carrion. The author has bred them from rotting bark
and decaying tea leaves. Almost any vegetable detritus which is moist and
not exposed to intensities of light or strong air currents seems to suffice. The
comparative rarity of these conditions at Crawley^ accounts for the small
numbers of Sciara sp. in the field. They breed in late fourth stage dungs-
more or less sheltered and practically devoid of other insects.

The larvae (PI, 1, Fig. 6) feed on dung which they comminute into a.
black powder only held together by moisture. This habit and their large
numbers render them a great obstacle to the successful rearing of Scarab,
larvae in the laboratory. There are intlications that they^ vill even attack
and consume Scarab pupae.

Pupation occurs in the dung v ithout formation of a pupal cell or cocoon.
The pupa has no respiratory processes, thougli these do occur in some species-
of Sciara ((.'sten Sacken, 1862). The j^uipal period \aries froii' two to five
days.

Copulation takes place at an early stage, sometimes before the wings of
the participants are properly iinfolde<l. Tlie ^\hole life cycle from egg to
adult required 10 to 12 days in June (average temperature, 14-5° C.) and
July (13*6° C.), 1941. In Denmark, Thomsen and Hammer (1936) estimated
the length of the life cycle of a species of Sciara to range from 19 to 22 days
at temperatures of 19° to 22° C.

The only other Sciarid foiuid at Crawley was a single specimen of ? Zygo-
netiro sp. bred from cow dung in the laboratory in July. 1941.

In spite of their short life cycle and fecundity, tlie Sciaridae are reiulered
of minor importance in the succession in cow dung by tlie frailty of the adults
and the restricted physiological requirements of the lar\ae.

F. 3. SCATOPSIDAE.

Tliese flies are also associated with a variety’’ of decaying materials.-
Scatopse sp. (? pidicaria) emerged from vessels containing pig, hors(', cow,
and calf dung in Denmark (Thomsen & Hammer, 1936), and a species occurs-
on carrion at Canberra.

A Consideration of the Insect Population Associated
WITH Cow Dung at Crawley, W.A.

233 :

At Crawley specimen of ? Rhegmoclema sp. emerged in large numbers
in the period November, 1940, to April, 1941, from dung brought into the
laboratory in November, 1940. In April, 1941, another batch came out of
39-day-old dung which also contained Scarab larvae. They have not been
seen in the field and consequently have no place in the succession in cow dung-
outdoors.

They appear to breed only in late fourth and such fifth stage dungs as
remain moist. Both the larva (PI. 1, Fig. 7) and the pupa (PL 1, Fig. 8) show
adaptations for living in poorly aerated, moist surrounding. Pupation occurs
inside the last larvae skin.

A number of adults dissected in February and March, 1941, contained
as many as six to eight nematodes, each in the abdominal region. The para-
sites must have occupied the greater part of the space of the abdomen.

F. 4. MYCETOPHILIDAE.

A single specimen of ? Mycetophila was obtained as a pupa from fifth
stage dung in the laboratory. Unlike the Sciarid pupa this was suspended
in a diffuse cocoon of white threads. The family is of no importance in the
succession.

F. 5, TIPULIDAE.

Two Eriopterine flies were obtained in July, 1941, from artificially-
moistened fifth stage cow dung. According to Imms (1938) the larvae may
occur in damp situations among grass, roots, decaying vegetation, or may
be aquatic. No sign of them in the field was seen.

F. 6. STRATIOMYIDAE.

While the adults of this family are mostly flower feeders, the larvae occur
in diverse habitata such as water, soil, and rotting wood (Imms, 1938). The
larvae of Myiochrysa have been found in cow dung (Williston, 1908). Actinia
incisuralis, Macq. is found near carrion (Fuller, 1934).

At Crawley, Neoexaireta spinigera, Wied. was seen to lay clutches of
eggs on fresh cow dung in the laboratory in May and June, 1941. These
hatched within three days, but the larvae failed to complete their develop-
ment. Other larvae collected on 7th November, 1940, which probably be-
longed to this species, reached the pupal period by 10th April, 1941, but no
adults emerged.

Stratiomyid larvae occur occasionally at Crawley in cow dung in the
field, although more numerous in rotting vegetation. These larvae, in the
course of their relatively long lives, must tolerate the change in condition
of the dung from fresh to old and dry, whereas in other dung-breeding insects
the larvae are generally restricted to the one stage or to stages which show
no marked differences.

F. 7. DOLICHOPODIDAE.

A number of black x)redaceous flies, near Asyyidetus sp., occur during
autumn and winter on fresh cow dung, preying on the Borborids attracted
to it.

F. 8. PHORIDAE.

Flies near Chaetocnemistoptera in Curran’s Key to the North American
genera (Curran, 1934) are frequently attracted to cow dung in the laboratory.
The relationship of carrion to cow dung insects holds good as Sciadocera
rufomaculata, WJiite, and Beckerma sp. are members of the carrion association
at Canberra (Fuller, 1934).

G. J. Snowball.

F. 9. SEPSIDAE.

Tliese are characteristic excrement insects with a world-wide distribution.
Sepsis viotacea, Meigen. breeds almost exclusively in excrement, including
that of man (Howard, 1912), There are 30 species of Sepsidae in Great Britain
(Imms, 1938). In Denmark, numbers of Sepsis sp. emerged from cow, horse,
and pig dung (Thomsen & Hammer, 1936). Sepsids are common in the
tropics on dung of all kinds (Patton & Cragg, 1913). In Australia “the
known species of the family occur in the adult stage on garbage, carrion, or
vegetation, some being very abundant on flowers. The larvae feed in manure
and carrion ” (Malloch, 1925A).

At Crawley, Sepsids are the most characteristic flies on cow dung. Two
species are present, viz., Australosepsis fulvescens. Mall., which appears to
contain only males, and its black variety atratula. Mall., of which both males
and females are known, and Sepsis pleheia, de Meij. Both species breed in
cow dung in the field.

A. fulvescejis is most numerous from November to May, and S. plebeia
during most of the year excej^t January and February. The latter species
was collected at Katanning in August, 1941. Both are members of the
carrion association at Canberra. A single specimen of S. hirsuta, de Meij.,
was bred from cow dung at Dog Swamp, Tuart Hill, W.A., in January, 1942.

The vSepsids appear on the fresh dung and remain until after the super-
ficial skin has become a firm crust, spending the time in feeding and oviposi-
tion. While parading on the dung they generally pair ofl and indulge in
remarkable antics. Each female carries a male, which applies its labella to
her cervical membrane and engages its legs on her prothorax. They remain
for hours thus, keeping up a ceaseless movement of legs and wings, yet tlie
time of actual sexual contact is short.

Oviposition has been observed only in Sepsis plebeia. The female, still
carrying the male, partly extrudes the egg several times before depositing
it with a sudden thrust of the abdomen. Usually buried in the dung, the
egg is occasionally merely dropped on the surface of the cake. Sepsids are
frecpiently seen ovipositing in dung in which eggs of large Muscids have al-
ready hatched. This is probably due to the Sepsids remaining for a
longer time on the dung, because it is their main food, than do the Muscids,
in the diet of wdiich it is only a subsidiary item.

Only one egg is usutilly laid in a particular spot. Constant returning of
the female to the same place results in the formation of large clutches of eggs
just below the surface with the interlacing respiratory processes (PI, 1,
Fig. 9) forming a white network which is often conspicuous on the surface.
The eggs hatcli in about 24 hours.

The larvae (PL 1, Fig. 10) are truly coprophagous, occurring mostly in
second and third stage dung, occasionally in the first and very rarely in the
fourth. The length of larval life is apjDroximately 10 days.

Pupation normally occurs in the dung itself, usually in the peripheral
layers, sometimes on the lower surface. The pupal period in September,
1941 (average' temperature, 13 9“ C.) was 23 days. Tlie total life history
from laying of the egg to emergence of th<^ adult took 33 days in September
aiid October (average temperature, 16*1° C.), 1941.

Because they occur in large numbers and not only consume but tunnel
through the dung, the la,rvae assist greatly in changing its nature. They are
parasitised at Oawley by an unidentified Alysiui which emerges from the
puparium. It is not an efficient pai'asite as infested stages are not common.

A Consideration of the Insect Population Associated
WITH Cow Dung at Crawley^ W.A.

235

F. 10. BORBORIDAE.

Malloch (1925B) mmtions the occurrence of larvae and adults of this
cosmopolitan family on manure and carrion in New South Wales. Two
species of Leptocera frequent carrion at Canberra (Fuller, 1934). In America,
Howard (1912) reared Limosina (= Leptocera) and Sphaerocera from cow
dung and human faeces. In India, Borhorus is often infected with a herpe-
tomonas as is Sphaerocera in France (Patton & Cragg, 1913). Thomsen &
Hammer (1936) bred Borborids in cow, calf, pig, and horse dung in Denmark.
They were present in large numbers on pig, sheep, and cow dung at Katanning
in August, 1941. At Crawley they are associated with carrion, rotting grass,
cow, and horse dung.

Most of the small black flies around cow dung at Crawley are Borborids,
species of Leptocera with a few others belonging to other unidentified Acalyp-
trate families. The largest species of Leptocera has actually been reared from
cow dung, though it is the least common in the field. The habits of the preg-
nant females of the smaller species leave little doubt that they also breed in
this medium.

The Borborids are abimdant on cow dung from May to October, with a
diminution in the complementary period. They rapidly appear on the fresh
dung. Unlike the Sepsids they can be found actually underneath the dung
pad. The adults feed on the fresh dung, the pregnant females becoming
remarkably distended. No information on the immature stages has been
gleaned at Crawley.

F. 11. DROSOPHILIDAE.

Though the usual association of Drosophilids is with fermentable materials,
some breed in cow dung. They are not common at Crawley in the field,
being most numerous in July, August, and September, during which period
their larvae may be found in second and third stage dungs. The Drosophilids
found on cow dung at Crawley and cow and pig dung at Katanning trace to
the genus Cladochaeta in Curran’s Key to the North American Genera.

The larvae are coprophagous and similar in habits to those of Sepsids.
Pupation occurs in the drier parts of the dung, the pupal period lasting about
three weeks.

F. 12. OTITIDAE.

Only two of the species of this family frequenting cow dung at Crawley,
Pogonortalis barbifera Macq. and Chrysomyza aenea, Fab., have been identified.
P. barbifera resembles the Sepsids in the fanning movements of the wings.
That C. aenea breeds in cow dung on occasion was shown by examination of a
sample of fourth stage cow dung from Armadale in March, 1941. This dung
had fermented to the extent that steam rose off it and it felt warm to the
touch, both unusual conditions. It contained a large number of puparia and
nearly mature larvae of C. aenea., pupae of Cercyon haemorrhoidalis, one of
Leptacinus socius, three larvae of Saprinus sp., and a pupa of Musca domestica.
Specimens in the collection of the Department of Agriculture, Perth, reared
from larvae taken in silage waste at Boyanup appear to belong to (7. aenea.

The larvae perform peculiar skipping evolutions when disturbed. The
body is flexed ventrally to enable the mouth hooks (PI. 1, Fig. 11) to grasp
the terminal part of the body, probably the raised area roimd the anus.
Then the longitudinal body muscles are strongly contracted. The mouth
hooks suddenly release their hold, producing sufficient force to flip the

G. J. Snowball.

2:U)

larvae up to a foot from its starting point. These acrobatics are similar to
those performed by the larvae of Piophila casei, the so-called “ Cheese
Skippers.” Continuous stimulation will cause a larva to repeat these move-
ments until it can no longer lift itself from the ground.

Pupation occurs in the dimg, the puparium being reddish-brown in colour.
ISTo exact figures of the length of life history are available. Larvae of imknown
age collected on 3rd April, 1941, had all given rise to adult flies by 30th May,
so that in some cases the life history requires more than 58 days, longer than
that of dung-breeding Muscids.

F. 13. CALLIPHORIDAE.

Sub-Family 1 : CALLIPHORINAE.

Calliphora (Neopollenia) australis, Boisd., Calliphora (Proeclcon) nociva.
Hardy, and Lucilia cuprina, Wied., all visit fresh cow dung in spring and au-
tumn, though never in large numbers. Attracted by the odour, the adults
suck up the liquid dung, often in large quantities, but no evidence of their
breeding in cow” dung at Crawley was obtained. Their association with it
is thus of a casual nature and can have no effect on the succession.

Sub-Family 2 ; SARCOPHAGINAE.

In the period October-Novembcr, 1941, 10 specimens of Sarcophaga (Para-
sarcophaga) depressa, Desvoidy, were reared in cow dung from Crawley, and
in January, 1942, 53 from dung obtained at Dog Swamp, W.A. The eggs are
deposited in fresh dung, especially in strong-stnelling cakes. In spite of
their large size, 20 to 30 larvae per cake may occur. They generally leave
the dung to pupate without displaying the same degree of restlessness prior
to pupation displayed by the larvae of Calliphora stygia. The cakes which
sheltered the larvae contained normal populations of other insects.

S. varia Walk, normally breeds in cow dung in New Zealand. Two species
of Sarcophaga frequent carrion at Canberra (Fuller, 1934), while various species
of the genus attack living sheep as secondary blowflies in the tropical and
sub-tropical parts of Northern Australia (C.S.I.R., 1933).

F. 14. MUSCIDAE.

Sub-Family 1 : FANNIINAE.

? Fannia ^p.

One fly probably belonging to this genus was bred from cow dung in the
laboratory at Crawley. The eggs were laid on 16th September, 1941, hatching
about eight days later into “hairy ” maggots. These remained on the sur-
face of the dung without penetrating it until 20th October, when four pupated.
The only fly which emerged at all did so on 2nd November.

In the field, eggs are common and larvae have been seen on two occasions,
having actually penetrated into third stage dung but no adults have been bred
out.

Sub-Family 2 : MUSCINAE,

Musca domestica, Linn.

Cow dung has very little attraction for the ovipositing female and this
fly has not been bred out of it at Crawley. A single puparium, wlaich subse-
quently gave rise to a female Musca domestica, was fjund in cow dimg from
Armadale which was of an unusual type, as described in the section on the
Otitidae, though it contained the usual insect population. The presence of
the puparium is remarkable, because the dung contained no admixture of any

A COKySIDEHATTOX OF THE IXSFCT POPULATIOX ASSOCIATED

WITH Cow Duxg at CkawleYj W.A.

237

other material. Where M. domestica is found breeding in cow dung, the
material is generally mixed with straw or some such substance (Austen, 1928.
Results quoted by Thomsen & Hammer).

Musca vetustissima. Walker.

This is the common bush fly which annoys workers in the open because
of its habit of clustering on the face and attacking the eyes. Its activities
have earned it the title of the Queensland cattle fly and it also pesters
horses. It is of some economic importance because it transmits Habronema
muscae, a Nematode parasite in the stomach of the horse and another unde-
termined species of Habronema which causes habronemic conjunctivitis of
horses (Johnston & Bancroft, 1920). It is probably capable of setting up
Anthrax infection under some circumstances (Cleland, 1913). The life history
has been described by Johnston &■ Bancroft (1919), the breeding medium
being either horse or cow dung. The total period of development from the
egg to the adult in the Eidsvald district of Queensland was found to be 10 to
14 days in November.

At Crawley the flies are numerous on fresh dungs although only six speci-
mens have actually been reared from dung in May, 1941, In January, 1942,
large numbers were reared from cow dung from Dog Swamp, Tuart Hill W.A.,
which also contained maggots of Sarcophaga {Parasarcophaga) depressa, Des-
voidy.

Sub-Family 3 ; ANTHOMYIINAE.

Hylemyia deceptiva, Mall.

A single specimen of this species or near to it was bred out from cow dung
in September, 1941. H. deceptiva is recorded on carrion at Canberra (Fuller,
1934). Another undetermined species of Hyl&myia was bred from second stage
cow dung in June, 1941.

H. variata Fallen and H. strigosa Fallen, which occur in Ayrshire and
Arran in West Scotland are ovoviviparous flies. Whether this is the case with
the local species is as yet unknown.

Sub-Family 4 • PHAONIINAE,

Rhynchomydaea (Hardyia) carinata, Stein.

Flies near this species are the commonest of the large Muscids frequenting
cow dung at Crawley. The adults, which feed on fresh dung, occur during the
period May to November, while specimens have been bred out from cow dung
in the period March to Noveml^er. The species is present at Katanning.

Eggs are laid singly ^vith the anterior ends projecting in fresh dung.
About 14 larvae is the maximum number wliich mature in each cake. They
occur in first to third stage dung, the larvae period being about 14 days. The
larvae are coprophagus.

Pupation occurs in the dung, usually in a hollowed cavity on the lower
or near the upper surface. Sometimes, after heavy rain, puparia may be
found on top of cakes in shaded situations. The pupal stage lasts about three
weeks. The total cycle from egg to emergence of the adult took 34 days in
September (average temperature 13*9° C.) and October (16*1° C.), 1941.

lO

G. J. Snowball.

:^S

HELINA.

This genus is wide-spread in Australia (Mailoch, 1925C) and is represented
at Crawley by the following species

Helina coerulescens, Sleie.

Though common on fresh dung at Crawley during the same period, this
metallic fly has not yet been reared from cow dung. It is similar in habits
to R. carinata.

Helina hypopleuralis, Mall.

Four specimens bred from cow dung at Crawley in September and No-
vember, 1941, belong to this species or to one near it.

Helina regina, Mall.

One specimen near this species was reared from cow^ dung at Crawley.

Muscina stabulans, Meig.

This wide-spread species has a variety of breeding media. Howard (1912),
in America, lists as food for its larvae, decaying animal matter, cow dung,
fungi, caterpillars, larval bees, dead pupae, decaying plant materials, and
human excreta. Fuller (1934) notes its association with carrion. In Denmark
it was reared from pig, cow, and calf dung (Thomsen & Hammer, 1936).
The flies have been reared from rotting potatoes and also from carrion
in Australia (Hardy, 1938), whereas Muirhead Thomson (1937) stated that
he had never seen nor had been able to induce oviposition by M. stabulans
on carrion.

Only one specimen has been reared from cow dung at Crawley.

Order 7— HYMENOPTERA.

F. 1. ALYSIIDAE.

An undetermined species of Alysiid is associated with cow dung at C'raw ley,
most numerous during the period February to November. In spite of their
delicate build they sometimes penetrate deeply into the interior of cakes,
presumably to parasitise Sepsid and Drosophilid larvae, since puparia so
infested are occasionally found. Fenales are more numerous than males.

An Alysiid, Alysia manducator Pantzer, has been introduced into Aus-
tralia as a parasite of sheep blowflies, but it has failed to establish itself (C.S.I.R.
1933).

F. 2. FIGITIDAE.

Small wasps belonging to this family are encountered throughout the
year in and on first stage dungs. As no material parasitised by them was
obtained, their hosts are not known. This is also true of a number of other
wasps belonging to various families, which were not identified because noth-
ing is known of their habits.

F. 3. FORMICIDAE.

On fresh dungs, small ants are usually present as predators attacking
adult Borborids and Sepsids. In due course they carry off immature stages of
beetles and flies. They are most active during the summer, at ^vhich time
they appear to influence the population considerably. Nevertheless, in
August, 1941, many of the second and third stage cow dcmgs were packed
with pupae which ants were removing.

A CONSIDERATIOK OB^ THE IXSECT POPTJLATIOX ASSOCIATED

'WITH Cow Dung at Crawley^ W.A.

239

Small amber-coloured ants frequently form colonies under dung cakes of
all stages in spite of the conclusions of Forel and Wheeler tliat they will not
do so except to seek shelter from the sun or moisture, both of which are amply
available at Crawley under stones or vegetation. Again, the odour does
not prevent ants from penetrating into even fresh dung, although Wheeler
(1926) refers to their dislike of bad odours such as those of carrion and mam-
malian faeces.

Order 8— LEPIDOPTEKA.

Three larvae collected on the 5th September, 1941, on top of late fifth
stage co-w dung about three months old, ]<ept in the laboratory. They were
apparently feeding on the dry and crumbling fibres of the cake. The three
larvae pupated five days later. After an additional 23 days, one gave rise
to a small Oecophorid moth. The larvae have not been seen in the field.

The association of these larvae with cow dung is of the same order as
Monopsis rusticella, Hubn., a Tineid moth breeding in the wool of an old sheep
carcase reported by Fuller (1934), being divorced in time and space from those
insects which form the succession proper.

8. OTHER ANIMALS ASSOCIATED MTTH COW DUNG AT CRAWLEY.

A. Other ARTHROPODA.

1. ACARINA.

A large number of mites belonging to the Parasitidae are found in cow
dung at Crawley. They breed in the dung and frequent all stages except the
final part of the fifth. A considerable -variation in the mite population of
indix idual cakes is noticeable.

In the field both flies and beetles are found carrying numbei’s of mites
which may merely hang by their claws or adhere by a secretion. Procto-
phanes sculpius, Hope, is especially liable to act as a carrier, as its clumsy build
and slowness of movement prevent it from dislodging its passengers.

Under laboratory conditions, where dung is kept covered, iuites multiply
to a remarkable extent. Though mites have not been seen, except in a few-
doubtful cases, to molest them, immature stages of flies and beetles diminish
in number in dungs w'ith a high Acarine population density.

At Katanning in August, 1941, no Parasitids were seen on cow dung in the
town itself and very few in material five miles away, a strong ntrast to the
large numbers present at Crawley in the same month.

2. CRUSTACEA.

Fourth and fairly moist fifth stage cow dung offe shelters associations
of Porcellio sp. at Crawley. They offer little or no competition to such im-
mature stages of Hydro23hilids or Scarabacids as are present.

B. NEMATOBA.

If kept in closed vessels, cow dung is an ideal nidus for these minute
worms. In the field they are not noticeable as a rule until pupae of Cercyon
begin to appear in fourth stage dung cakes. A number of these have bunches
of Nematodes hanging from the appendages or on the ventral body surface,
often numerous enough to force the wings of the j^ujDa into an unnatural |)osi-

240

Cr . J. Snowball.

tion almost at right angles to the long axis of the body. Since most of the
pupae in this condition are able to transform normally, it seems that the
Nematodes are free living forms attracted to them by the moisture given off.

Adult Cercyons, when kept in closed vessels with cow dung, often con-
tain Nematodes in the posterior part of the alimentary canal and genital tract
as well as on the moist body surfaces. There is no reason to believe that the
presence inside the body is due to anything more than the tendency to migrate
into any aperture.

C. ANNELIDA.

Fresh cow dung attracts earthworms which will consume large quantities
if it is applied to garden beds containing them. They are also of frequent
occurrence in heaps of fourth stage cow dung which has retained moisture.
Nothing definite is known of the degree of competition they offer to the true
dung insects.

9. FEATURES OF THE SUCCESSION IN COW DUNG.

Having referred to various aspects of tlie biology of the insects concerned,
it is now^ possible to draw up a scheme showing the features of the succession,
defined according to the stage through which the cow dung passes, as set out
in section 4.

First Stage.

This is characterised by the activities of dung insects on the surface of the
fresh dung. The Sepsids, Borborids, Calliphorids, Otitids, and Muscids
appear to feed on the liquid dung in such a haphazard manner that the order
varies for practically every dung.

The dung bettles are also prompt in arrival and again no particular order
can be defined. The Staphylinids, Hydrophilids, Histerids (in season) and
Aphodiines, except A. ambigims, all commence to penetrate either through
natural crevices on both surfaces or by creating apertures in the superficial
skin.

Toward the end of the first stage, the number of flies diminishes. The
Calliphorids leave first, the only attraction for them being the soft dung now
covered with a firm skin, and then the large dung-breeding Muscids after ovi-
position. The Sepsids and Borborids remain longer, and the former may be
seen ovipositing over some days. The predatory Dolichopodids and Creo-
philus erythrocephalus also leave when the numbers of their prey diminishes.

Second Stage.

The surface layers of the cake become penetrated by the dung beetles.
Sepsids and Borborids are still present. Mites and Collemi^ola appear early
in this or late in the first stage. Although the eggs of Muscids and Sepsids have
hatched, the small size of the larvae renders them inconspicuous. Adult
Psychodids are to be found o\'ipositing on cakes in the shade.

Third Stage.

The dung is extensively penetrated by beetles. In many cases an un-
jienetrated core near the low^er surface is left but many cakes are completely
riddled. The activities of Cercyons and Aphodius lividus, by admitting air
into the dung, give it its characteristic black colour. The latter part of this
period is primarily one of immature stages. The poi^ulation of Sepsid, Droso-
philid, and Psychodid larvae becomes very high. Cercyon and even occasional
Scarab larvae aj^pear tow^ards its end, as the emigration of adult beetles begins.

A CONSIDKHATIOX OF TIJE IXSFOT POFULATIOX AsROCTATKD
WITH Cow DUXO AT (-RAAVEHY, W.A.

241

Fourth Stage,

Except for occasional specimens, usually Staphylinicis, the population of
adult beetles is low apart from Aphodius amhiguus. Larvae and pupae of
Staphylinids, Cercyons, eggs and larvae of Aphodiines are present in the peri-
pheral regions. The Psychodids have disappeared. Larvae of Sciara sp.
hatch out in tliis stage. Most of the Brachycerous Dipteran larvae have
pupated. Mites and Collembola still remain in some numbers.

Fifth Stage.

The population is essentially similar to that of the previous stage, progres-
sively diminishing in size as the dung becomes drier. Ultimately, in the
final stages, the Aphodiine larvae have undergone metamorphosis and vanished.
The dung no longer contains insects and serves only to shelter a few, the
presence of which under it is purely fortuitous.

The following minor successions compose the main one : —

1. The succession of visitors to inhabitants, which arises out of the

alteration of the dmig, due more to oxidation and other chemical
changes at its surface than to any activities of the insects.

2. The succession among the adult Coleoptera. The majority of the

early invaders move out in the third stage and are succeeded by
Aphodius amhiguus in the early fourth, which is replaced in
turn only by the casual shclterers of the fifth. The cause of
the emigration in the third stage is not always clear. High
population densities may cause a rapid change in the quality
of the dung which becomes unsuitable for the adults and con-
sequently they leave. There are, however, many instances
in which the insects abandon the dung v’hile it still offers abund-
ant food and space. Competition may cause the emigration of
surplus individuals. This would lessen its intensity and there
would be no reason for any more to leave. The exodus of
all beetles from a dung of this type may be due to the tendencies
found among gregarious insects to keep close together and to
imitate each other’s movements, which are believed to be re-
sponsible for such phenomena as the swarming of locusts (Uvarov,
1928). It is an open question whether the gregariousness
of dang beetles is real or due to purely chance associations
resulting from largo numbers of individuals responding to the
same stimulus. In any case, the migration of dung beetles is
a gradual jorocess not comparable with any form of swarming.

3. The succession among the larval stages. Larvae appear in cow

dung in the following order : —

1st Stage — Sepsid, Drosophilid, Borborid (?), Miiscid.

2nd Stage — Psychodid.

3rd Stage — Hydrophilid, Staphylinid.

4th Stage — Aphodiine, Sciarid.

In carrion the succession is assisted by the predaceous activities of the
secondary blowfly maggots which actually attack or drive off those of the
primaries. In the cow dung i)opulation none of the larvae of importance in the
succession arc predatory on the others except those of Staphylinids, which do
not appear in sufficient numbers to influence the total, Aphodiine larvae
succeed those of Cercyon only because their physiology is better adapted to
existence in fourth stage dung. The effect of competition is therefore to
regulate the number of individuals found in any stage and plays little part
in replacing one species by another.

G. J. Snowball.

-4‘>

Chapman (1931) divides animal successions into passive and active
according to the part played by the animals concerned. Though less so than
that in carrion, the succession in cow dung is of the active type since the
insects do affect their environment. Volume for volume cow dung supports
a smaller poj3ulation tlian carrion. This is due to the less powerful odour
which is the primary attracting stimulus, the less nutritious nature of the
dung itself , and in the case of flies the comparatively short time during which
it is a suitable medium for oviposition owing to the early formation of a more
or less impervious crust. All these factors result in a less well-defined succes-
sion tlian occurs in carrion.

As has been indicated previously, there is a correlation between the type
of insect in carrion and that in dung, many species being common to both
habitats. Nevertheless, each substance has its own characteristic insects
physiologically adapted to it with only a casual association with the other.
Jllowflies visit cow dung but their life history is bound up with carrion. In
the same way, Cercyons are sometimes found on carrion yet they are character-
istic cow dung insects.

Both in carrion and cow dung are insects which either have a wide dis-
tribution themselves, e.gr., Aphodius lividus, Cercyon haemorrhoidalis, or
have closely allied species in other parts of the world, e.g.. Sepsis plebeia
This is to be expected, as neither carrion nor faecal matter is subject to geo-
graphical variation.

10. THE ECONOMIC IMPORTANCE OF INSECTS FOUND IN COW

DUNG.

The primary sheep blowflies Callipliora australis Boisd, C. nociva Hardy,
and Lucilia cuprina Wied., have only a casual association with various species
of Sarcophaga, are secondary sheep blowflies, and some of these breed in cow
dung, though the importance of this material to the flies as a breeding sub-
stance is not known. Musca domestica Linn, prefers other breeding nidi,
and no evidence has been seen that Stomoxys calcitrans Linn., which occurs
in numbers in Perth, ever frequents dung at Crawley. Research in various
parts of the world has showm that cow dung is among the least favoured of
the breeding media of this fly.

Musca vetustissima Walker is a common pest in Perth on man and cattle.
Cow dung is an important source of the fly here, especially that dropped in
swampy situations. Rhynchoniydaea carmata Stein and the species of Helina
associated with cow dung at Crawley are not domestic insects, nor have they,
as far as is known, any habits rendering them objectionable to man. Muscina
stabidans Fallen is not abundant at Crawley. In any event, cow dung is
only one of a large number of breeding materials utilised by this fly.

In South Australia and Tasmania, two Aphodiines, which normally feed
on dung, have come under notice because their lavae have been found damag-
ing pastures and lawns (Swan, 1934 ; Evans, 1941). These are Aphodius
howittiy Hope, and A. tasynaniae^ Hope, neither of which has been collected at
Crawley. No complaints have been received by the Department of Agri-
culture against any local species in this connection.

When L. J. Newman, in 1929, succeeded in rearing Lyperosia exigua de
Mcij., the Buffalo fly on cow dung under midwinter conditions in Perth, with-
out the application of artificial heat, it was feared that this pest might be
introduced in the dairying regions in the southern part of the State. Pre-
cautions against this possibility in the form of sprajdng cattle before they
left North-West jiorts were introduced (Tcoj^, 1931). The fly has not been
reported in the South-West, nor has any sign of it been seen at Crawley.

A Consii)p:ratiox of tup: Insect Population Associated
WITH Cow Dung at Crawley, W.A.

243

The typical dung insects — Sepsids, Hydroplhlids, Aphodiines, etc., all
])lay their part as scavengers in getting I’id of wast(‘ material, but the actual
evaluation of the economic importance of tliis ])art is practically impossible.

AC^KNOWLEDGMENTS.

The writer wishes to express his indebtedness and gratitude to the follow-
ing, while absolving them from any responsibility regarding statements of
opinion by the writer : — Professor O. E. Nicholls for general lielp and guid-
ance ; Dr, A. J. Nicholson for suggesting the nature of the problem studied ;
Mr. K. C. McKeown and officials of the Australian Museum, Sydney, for the
identification of the Coleoptera ; Mr. G. H. Hardy for the identification of
most of the Diptera ; Mr. J, H. Perkins for the identification of Sepsidae
and Borboridae ; tlie Officers of tlie Entomological Ih'anch of the W.A. De-
partment of Agriculture for their co-operation ; Dr. G. A. Currie and Mr.
K. R. Norris for advice on various aspects of the problem.

REEERENCPiS.

Austen, E. E. (1928). — The House Fly, 3rd. ed., Brit. Mus. Bcon., Series No. lA. (Reference
not seen, quoted by Thomsen & Hammer, 1936).

Blackburn, Rev. T. (1904). — Proc. Roy. Soc. Vic., Vol. XVII. (New Series), Part 1, Sept.,
1904.

Boucomont, A. (1929). — Linguan Science Journal, Vol. 7, June, 1929. “ A List of the

Coprophagous Coleoptera of China.”

Broun, T. (1880). — Manual of N.Z. Coleoptera, Parts 1-7, 1880-1893 {Colonial Museum
and Geol. Survey Dept.). Parts 1-4 [N.Z. Institute, Parts 5-7).

Chapman, R. N. (1931). — Animal Ficology (McGraw Hill).

Cleland, J. B. (1913). — “ Insects and their relationships to disease in Man in Australia.”

Trans. Aust. Med. Congress (Sydney, 1911), 9, 1913, pp. 548-570 (Reference not seen, quoted,
by Johnston & Bancroft, 1920).

C.S.I.R. (1933). — Pamphlet 37 and N.S.W. Dept, of Ag. Sc. Bulletin No. 40, “ The Sheep
Blowfly Problem in Australia.” Report No. 1 by Joint Blowfly Committee.

Curran, G. H. (1934). — The Families and Genera of North American Diptera.

Davidson, J. (1937). — Austr. Jour. Exp. Bio. and Med. Sci., Vol. 15, The Temperature-
Developmental Curve of L. exigua in relation to its probable distribution in Australia.

Esdaile, P. (1927). — Economic Biology for Students of Social Science. Part 1 — Harmful
and Useful Animals.

Evans, J. W. (1941). — Tas. Jotirn. of Agric., Feb., 1941 ; De])t. of Agric., Vol. XII. (New
Series), No. 1.

Fuller, M. E. (1934). — C.S.I.R. Bulletin No. 82, “The Insect Inhabitants of Carrion, A
Study in Animal Ecology.”

Graham, S. A. (1925). — The Felled Tree Trunk as an Ecological Unit. Ecology 16 : 347
(Reference not seen, quoted by Fuller, 1934).

Hardy, G. H. (1938). — Proc. Roy. Soc. QM., Vol. L., “ Notes on Australian Muscoidea IV.”

Howard, L. O. (1912). — The House Fly, Disease Carrier.

Imms, A. D. (1938). — A General Textbook of Entomology.

Johnston & Bancroft (1919). — Proc. Roy. Soc. Qld., Vol. XXXI, 1919 (20), pp. 181-203,
“ Life History of Musca australis and M. vetustissima.

Johnston & Bancroft (1920). — Proc. Roy. Soc. Qld., Vol. XXXIl., 1920. The Life History
Habronema in relation to M. domestica and native species in Queensland.

Lacordaire, T. (1854). — Histoire Naturelle, Genera des Coleopteres. Tome Premier.

Lea, A. M. (1923A). — Reeds. S. Aus. Museum, Vol. 11, No. 3, 1923. Australian Dung
Beetles of Sub. F. (-oprides (four plates).

Lea, A. M. (1923B). — Proc. Roy. Soc. Pic., Vol. XXVI. (New Series), Part 1, Sept., 1904.

Lefroy, M. (1923). — Manual of Entomology.

Lindquist, A. (1935). — Circular 351, U.S. Dept, of Agric. “ Notes on Coprophagous
beetles and methods of rearing them.”

Mackerass, I. M. (1932). — C.S.I.R. Journ., 1932, Vol. 5, No. 4. “ A note on the occurrence

of Hydrotea australis in N. Aus.”

Malloch, j. R. (1925A). — Proc. Linn. Soc. N.S.W. , Vol. 50, Part IV. “ Notes on Australian.
Diptera, Part VII.”

Malloch, J. R. (1925B). — Proc. Linn. Soc. APiSMF., Vol. 50, Part II. “ Notes on Australian
Diptera, Part VI.”

244

G. J. Snowball.

Malloch, J. R. (1925C).~P/-or. Linn. Hoc. .V..S'jr., Vol. 50. Part 2. “ Notes on Australian

Diptera, Part V.”

Myers, J. G. {1938}.— Jowr«. C.H.I.R., Feb., 1938. “ Report on Natural Enemies of the

Horn Fly (Lyperosia irritans) in Haiti.’'

Muirhead Thomson, R. 0. (1937).— Para, sMor/y, Vol. XXIX., No. 3. “Observations on
the Biology and Larvae of the Anthomyidae.”

Osten Sacken. C. R. (1862).— Pror. Fnt. Hoc. Pliiladelphia, 1862. “ Characters of the Larvae

of the Mycetophilidae.”

Patton & Cragg (1913). — Textbook of ^Medical Entomology.

Schmidt, Adolf (1922).— Has Tierreich, Coleoptera, Aphodiinae.

Sharp, H. (1884). — Trans. Ent. Hoc. London, 1884. “The 'Water Beetles of Japan ” pp.
439-464. “ Revision of Hydrophilidae of N.Z.,” pp. 465-480.

Sharp, H. (1890). Trans. Ent. Hoc. Lond., 1890. “On some aquatic Coleoptera of Japan”

Simmons, P., Reed, W. H., McGregor, E. A. {\m\).—U.H. Dept, o fAqric. Circ. 157, April
1931. ■■ Fig Insects in C'alifornia.”

Skuse, F. A. A. (1889).— Pror. Linn. Hoc. N.H.W., Vol. 3. “ Hiptera of Aus., Part 2— Sciari-

dae.”

Spry, W^, and Shuckard, •\\\ (1840).— The British Coleoptera Delineated.

Stephens, J. F. (1839). — A Manual of British Coleoptera.

Swan D. C. (1934).— Jowtk. of Agric. of H. Aus., Vol. 37, 1149-1156. “ A Scarab Beetle

(Aphodius tasmaniae Hope) destructive to ])astures in S.E. of S. Australia.”

Thomsen & Hammer (1936). — Bull. Ent. Research, XX\TI. “ The breeding media of some
common flies.”

Tillyard, R. J. (1926).— The Insects of Australia and N.Z.

Toop, C. R. (1933).— Measures for the Prevention of Spread of the Buffalo Fly. Journ.
Dept. Ague. W .A., Dec., 1931, Xo\. 8 (2nd Series), No. 4.

Dvarov, B. P. (1928). — Locusts and Grasshoppers.

Warburton, C. (1909). — Cambridge Natural History, Vol. IV. Crustacea and Arachnids.

Wheeler, W. M. (1922). — Social Life of Insects.

Wheeler, W. M. (1926). — Ants.

Williston, S. M. (1908). — Manual of North American Diptera.

EXPLANATION OF PLATE 1.

Fig. 1. — Cercyon liaemorrhoidalis Fab. Variety 1.

Ventral view of Adult.

Fig. 2. — Cercyon haemorrhoidalis Fab.

Dorsal view of Mature Larva.

Fig. 3. — Lepiacinus socius Fauv.

Lateral view of Pupa.

Fig. 4. — Aphodius lividus Oliv.

Ventral view of Pupa.

Fig. 5.. — Psychoda sp.

^’^entral view of Larva.

Fig. 6. — Hciara sp.

Partially dorsal view of Larva.

Fig. 7. — ? RJiegmoclema sp.

Lateral view of Larva.

Fig. 8. — ? Rhegmoclema sp.

Lateral view of Pupa extracted from skin of last larval instar.

Fig. 9. — Sepsis plebeia, de Meij.

Kgg.

Fig. 10. — Sepsis plebeia, de Meij.

Lateral view of terminal segments of Larva.

Fig. 11. — Chrysomyza aenea Fab.

Lateral view of anterior region of nearly mature Maggot.

Fig. 12. — Rhynchomydaea (Hardyia) carinata Stein or near.

Posterior spiracles of second Larva.

A CONSIDERATIOX OF THE INSECT POPULATION ASSOCIATED

AviTH Cow Dung at Crawley, W.A.

245

4

The Essp:ntial Oils of the Westehn Australian Eucalvpts. 247

13.— THE ESSENTIAL OILS OF THE WESTERN
AUSTRALIAN EUCALYPTS.

Part VII.

THE OIL OF EUCALYPTUS EBYTHRONEMA, TURCZ.

Bv E. M. Watson, Ph. D.

Read 9th June, 1942 ; published 28th March, 1944.

EiicoJyptus crythronerna Turez. has been recorded as occurring along the
Eastern Railway from Tammin to Southern Cross. North of Tammin, it is
known at Wyalkatchem and Cow'cowing and, farther west and north-west,
at Wongan Hills and Ballidu. It also occurs soutli of the railway, and the
material used in this investigation was collected some 10 miles south-east
of Merredin.

It is a small tree, 12 to 18 feet high, or a malice with a bulbous stock
and several erect stems. It grows in clay soil, usually associated -with E.
saliihris and E. salmonophloia^ either in the malleo association fringing the
woodland or in the woodland. The leaves are lanceolate, narrow^ lanceolate,,
or falcate in shape, and are from tw'O inches to four inches long. The colour is a.
dark and usually lustrous green, the venation of the cineole type and the-
submarginal vein is removed from the edge. The lamina is copiously dotted,
with oil glands. The bark is covered wdth a fine white talc-like powder which
easily rubs off, leaving, in the older parts, a pinkish bark. Because of this;
white powder on the stem and branches, it is commonly called the ‘‘ white
mallee.” The wood is dark brown in colour, is very hard and susceptible to-
termite attack, which, how'ever, is confined to the heart wood.

The material used was collected by Mr. G. E. Brockw-ay of the ForestS'
Department, towards the end of August, 1941. It was obtained from mallees
on Avon Location 20751, about 10 miles south-east of Merredin, and its identity
was confirmed by Mr. C. A. Gardner, Government Botanist.

The air-dried terminal branches were steam distilled, the oil coming'
over very rapidly. It w^as pale yellow in colour and had an irritating odour
the yield a\’craged from 2*5 to 2-6 per cent, by weight. The physical con-
stants of the oil are those of a typical cineole oil, and its solubility in alcohol
shows a low proportion of terpenes. The crude oil itself contains over 70
per cent, of cineole. whilst the rectified oil contains 80 per cent, of cineole.
Free acids and esters are present in only small amount, but alcohols, the greater-
part of which is geraniol, make up more than 13 per cent, of the oil. Both,
low' and high boiling aldeliydes are present, but in no great quantity.

The rectified oil, boiling between 164° and 185°, is a very high grade-
medicinal oil. It makes up over 80 per cent, of the oil, contains, as already
stated, 80 per cent, of cineole, and (‘omplies wdth all the requirements of the
British Pharmacopoeia for eucalyptus oil. It is colourless and has a very
pleasant, non-irritating, camphoraceous odour.

Although clearing for farming has probably removed appreciable
quantities of this eucalypt, its commercial exploitation should still be possible..
Cultivation shoukl also be economically sound on account of the good yield
of high grade oil obtainable from it.

248

E. M. Watson.

EXPERIMENTAL.

The oil distilled completely in three hours and the undried oil had the
following pljysical properties at 20° (.!. : — Specific gravity, 0-923 ; refractive
index, 1-406: specific rotation, -0-22°. Tt ^cas soluble in two volumes of 70
per cent, alcohol. Its acid A-aluc was 0-6 : its saponification values Avere : —
Cold, 1-9 ; hot, 3-2, corresponding respectively to 0-07 per cent, of geran^ 1
acetate and 1 ■ 1 per cent, of total esters calculated as 012 ^ 20 ^ 2 - saponifi-
cation values of the acetylat.ed oil \Aere : — Cold, 43-8 ; hot, 52-5, correspond-
ing respectiA^ely to l]-r> per ceTit. of alcohols calciilated as geraniol and 13-5
per cent, of total alcohols calculated as CioHigO. The aldehyde content of
the oil A\as 0-12 millimole per giam of oil, ec|iuA'alent to 1-8 ])er cent, of
aldehydes calciilated as CJ 0 H 14 O. Tlio cineole content of the dried oil was
70-9 per cent. The usual colour reaetious for aromadendrene Avere obtained
but the oil gave no reactions for ])inene or pliellandrene.

The oil Avas redistilled and tlie follow ing fractions Avere separated : —

Fraction.

Boiling

Amount

Specific

Re tract iA’e

Specific

Range.

per cent.

GraA’ity.

Index.

Rotation.

1

tqi to 160°

2-8

0-897

1-442

714-1°

2

16C°— 167°

11-75

0-901

1-462

+ 16-1°

3

167°—-169°

11 -25

0-912

1-461

+ 6-5°

4

169°— 176°

57-4

0-920

1-461

1-2°

The

folloAAing

residue was further
fractions separated

fi’actionated

under

reduced pressure

and til

5

80°— 90° /27 mmy.

2-2

0-959

1-480

—23-8°

6

90°— 110°/26mms.

6-0

0-967

1-494

—36-2°

7

110°— 120°/25 mms.

1-7

0-965

1-496

—19-7°

As the initial boiling commenced, the formation of A\ hite insoluble material
w^as noted and from the final residue 2-3 per cent, of AA'hite amorphous solid
AA^as separated by addition of ether, folloAAed by filtration.

Fraction 1 contained some Avater which was separated. This contained
both aldehydes and free acids, the latter including butyric, probably A^aleric,
and either or both formic and acetic acids. From this aqueous portion a
small quantity of colourless crystals se]iarated. These crystals are neutral,
soluble in alcohol, may be recrystallised from Avater, give no colour Avith
ferric chloride, and melt at 145°. From the oily part of this fraction no re-
actions for pineno were obtained.

I'raction 2 contained 59-3 per cent, of cineole, A\ as colourless and pleasant-
smelling, and bad cold and hot saponification A'alucs of 5-9 and 8-2 rcspect-
iA^eJy.

Fraction 3 was similar and contained 76-5 pei’ cent, of cineole, Avhilst
Fraction 4 contained no less than 88-8 per cent, of cineole. The saponifica-
tion values of these two fractions were respectively 2-2 and 2-9 (cold) and 3- 1
and 3-6 (liot). No crystalline nitrosito aars obtained from Fractions 2, 3,
and 4.

I’raction 5 was colourless and contained the greater part of tlie esters,
haAnng cold and hot saponification A’alues of 56-6 and 63-9 respectively,
corresponding to 20-8 and 22*4 per cent, of esters, calculated as Ci 2 R 20 ^ 2 -
This fraction, as well as fractions 6 and 7, contained a small amount of
aldehyde.

Fractions 6 and 7 had cold saponification valaes of 19- 1 and 15-8 respect-
ively and hot saponification values of 22-5 and 20-3.

The Essential Oils of the Western Australian Eucalypts. 249

Fraction 7 (wliicli was pale yellow in colour) and the final residue both
.gave a purple colouration with ferric chloride.

Distillation of 150 grams of the oil gave 123 grams (82 per cent.) of oil,
distilling between 164° and 185°. This oil contained 80 per cent, of cineole
and had the following physical properties at 20° ; — Specific gravity, 0*918 ;
refractive index, 1*461; specific rotation, -!-2*76°. It was soluble in two
volumes of 70 per cent, alcohol. The requirements of the British Pharma-
copoeia for eucalyptus oil are : — Cineole, not less than 70 per cent. ; specific
gravity (15*5°/15*5°), 0*910 to 0*930; refractive index (20°), 1*458 to
1 *470 ; optical rotation, — 5° to +5° ; soluble in five volumes of 70 per cent,
alcohol. The oil gave no reaction for phellandi'ene and, when tested for
aldehydes by the Pharmacopoeial method, required 0*65 ml. of 0*5N alcoholic
caustic alkali (B.P. fi.gure, not more than 2 ml.).

The author is indebted to Mr. G. E. Brockway for the collection, and to
Mr. C. A. Gardner for the identification of the material.

Perth Technical College.

General Index

Ixxxix.

GENERAL INDEX.

Amphibolites, of Toodyay district
Amphibolites, analyses
Andalusite, Toodyay district
Arthur River
Aulosteges

Aulosteges baracoodensis
Aulosteges medlicottiarms
Aulosteges spinosus ...

Aulosteges tibeticus ...

Aulosteges wangenheimi
Aulosteges wolfcampensis
Avonia

Page

... 104, 111, 119

105

101, 129

13

11, 12, 32, 39, 40
7, 29, 32
... 32, 35, 30

30

32, 35

S. 32, 33, 34

32

30

Ballythanna Hill
Balmaningarra
Barrabiddy
Barragooda Pool
Booker, F.
Branxtonia
Branxtonia typica
Broili. F.

38, 40,
i'o, 17, "l8, 20, 23,

mom, r. ... ... ... ••• ••• ••• ••• ^ *•» ’

Bryozoa from the Upper Palaeozoic of Western Australia. A Revision of some pre-
viously described species of

Bryozoa from the Wandagee and Nooncanbah Series (Permian) of Western Australia
Bullsbrook, fossil plants from

27

54

54

13

40

41
40
20

187

105

201

Calceola

Calc-silicate rocks, of Toodyay district

Callytharra Springs ...

Cancrinella
Capertree, N.S.W.

Centropages australiensis, sp. nov.

Chao, V. T

Chapman, F. ...

Chidlows
Chonetella
Chonetes
Chonetinae
Chonopectus ...

{'hristmas Creek Homestead
Chrome-muscovite, in Toodyay (piartzites
Cladophlebis australis
Climate of Western Australia
Clothoda nobilis

Cockburn Sound, Pelagic copejmds from

Condra, G. F.

Coniopteris hymenophylloides
(^onoclypiis in North-West Australia

Conoclypus westraliensis, .sp. nov.

Coolkilya Pool
(’ooper, G. A.

(bpepods, Marine from Western Australia
Copiapite

Cordierite anthophyllite rocks

C'ow Dung, Insect po]>ulation associated with at (,-rawley
Crawley, Insect population associated with cow dung at

9

94, 118
1,3, 21, 30, 43

32

42

209

... 10, 17, 20, 23, 25

13, 21, 22. 25

79

10

... 9. 12

12

10. 11, 12, 53

50

91

201, 203

XXX.

140

209

8. 11. 40, 53

203

75

75

27, 47

4, 13

209

79

118

219

219

xe.

General Index.

Cretaceous, Lower, plants ...
Gummingtonite, in schist, Toodyay district

Page

201

100

Davidson, T

Daviesella
Diciyoclostuft ...

Dictyoclostus callytharrensis, sp. nov.
Dictyoclostus callytharrensis var. wadei
Dictyoclostus gratiosus
Dictyoclostus indicus
Dictyoclostus semireiiculatus
Dictyoclostus spiralis
Diener, C.

Dolerite, quartz of Toodyay district
Dolerite, quartz, analysis ...

Douglas, J. A.

Dun, W. S

Dunbar, C. 0.

Dung, Insects associated with cow ...

... 30, 31, 45

10 , 12

9, 12

4, 13, 19

16, 19

17

4

29

18

11, 18, 20, 21, 23

125

127

23

21. 27

8, 11, 40, 53
219

Echinoidea

Eighty-mile beach. Ecological notes on the vegetation of

Elatocladus

Elatocladus pla-na

Embioptera of Western Australia, Revision of
Eragrostis epipoda

Etheridge, jr. R 13, 21, 22, 23, 27, 32, 39, 40, 42, 43,

Etheridgina

Etheridgina muirwoodae, sp. nov.

Essential Oils of the Western Australian Eucalypts. Part \’TI.
Eiwalyptus erythronema, oil of
Eulvsites, rocks related to

5, 46

47,

75
157
201
204
139
155
54, 56
40. 52
54

... 247

... 247

113, 115

10,

Fenestrellina affiuensa

Fenestrellina cacuminatis, sp. nov.
Fenestrellina columnaris, sp. nov.

Fenestrellina disjecta, sp. nov

Fenestrellina horologia

Fenestrellina lennardi, sp. nov

Fenestrellina ruidacarinata, sp. nov.

Fenestrellina valentis, sp. nov

Fibroferrite and Copiapite from Yetai' Spring.

Foord, A. H.

Fossil Cliff

Fredericks. G.

near C

lidlows

9,' 13, 2

188

171

170

168

167, 189

171

169

169

79

22,47

27, 43. 46, 51, 52
2, 5, 29. 35

Gascoyne River

Gedrite, in schist. Toodyay district
(rigantella

Gingin. fossil plants from ...

Girty, H

Glass Mts., Texas
Glauert, L.

Gnarrea Pool

Gneiies, granitic, analyses ...

Gneiss, granitic, of Toodyay district
Gold, in Toodyay ilistrict ...

(b’anite, and related rocks of Toodyay district
Gray, J. E. ...

Greger. I). K.

Grossularite, Toodyay
Grill lerite. Toodyay ...

22

!!! 100

12

201-7

32

8

13, 21. 22, 27

24

no. 123
107. 114

129

124

9

53

95

117

Oenkral Index

xo\.

Hamlet, B. ...

Helmersen, G.

Heteralosia

Heteralosia slocomi ...

Hill, C

Hornblende schist, Toodyay district
Hornblende schist, analyses
Hosking, L. V.

Page

17, 18

32

12, 40, 41

41

47. 50

104, 111

105

13. 15, 22, 27, 30. 36. 37

Insect popnlation associated with Cow Hung at Crawley, W.A., a consideration of ...
Isoetes ...

Isoetes Boryana
Isoetes Engelmanni ...

Isoetites

Isoetites Chaffatl

Isoetites elegans, sp.

Isoetites horridus
Isoetites serratus

219

201

202

202

201

202

202

202

202

Jack, L. K. ...

.Hmba Jimba Station

Jimperding Series, at Toodyay, petrology of
Jurassic plants from Gingin

21, 27, 42. 43
... 13, 42, 47

88

201-7

King, R. E. ...

King, W

Koninck, L. de
Kozlowski, K.
Krotovia

Krotovia micracaniha
Krotovia senticosa
Krotovia spimdosa

8

9, 37, 45
... 21, 42
20, 23. 54

29

30

30

30

Lepidocyclina ...

Leptalosia
Le Vene, C. ...

Eichaiew, B. ...

Linoprgductus ...

Linoproductus cancriniformis
lAnoprod'octus cancriniformis var. lyovi
Linoprod'octns cor a ...

Linoproductus cora var. foordi
Eniluigiii

Lyropora erkosoidcs ...

76

12. 40, 53. 54

29

11, 12, 34, 35, 39

7, 20

23

24

20

22

25

191

Machairopus cockburni, sp. nov

Marginifera ...

Marine Copepods from Western Anstialia, Series 11. ...

Martin, W

Metoligotoma ...

Middalya Station

Minilya, gen. nov

Minilya duplaris, s]). nov

Minilya princeps, sp. nov

Minilya River
Morris, .1.

Mount Abbott
Mount Cedric
Mount Dromedary
Mount Hardman
Mount Marmion

17, 24. 3

4 27

210

18

209

13

141

13

172

173

174
47. 54

27
47
26
27
25
54

XCll.

General Index.

Mount Nicholson
Mount Wellington
Muir-Wood, H.

0, 7, 8,

Page
16
27
12, 31

Netschajew, A.

Nilssonia
Nooncanbali ...

Nooncanbah Series (Permian) of Western Am
North-West Basin
Notoligotoma ...

Notoligotoma hardy i ...

Notoligotomidae

Oehlei't, D. P.

Oil of Eucalyptus erythronema
Oligotoma

Oligotoma approximans
Oligotoma glauerti
Oligotoma gurney i
Oligotoma gurneye spinulosa
Oligotoma tillyardi
Oligotomatidae
Orbigny, A. d'

Orthis ...

Orthotetinae ...

34

201

47

l alia. Bi vozoa from the Wandagee and 165

... ‘ 9

144

144

144

52, 53. 54
... 247

141

142

141

... 143

143

142
... 141

20

9

3

Paeckelmann, W.

Pecten ...

Pelagic Copepods from Cockburn Sound
Phyllopteris

Polypora fovea, sp. nov

Polypora multiporifera, sp. nov

Polypora relificis, sp. nov

Polypora, sp. nov. indet. ...

Polypora ^voodsi
Prendergast, K. L. ...

Proboscidella ...

Productella
Productellinae
Productina
Productinae ...

Productorthis ...

Productus complertems
Prod actus cora
Productus corrugatus...

Productus latirostrus ...

Productus line.aius
Productus neffedieri ...

Productus ocatus
Productus prafteniauus

Productus semireticulatus {fee also Dictyoclostus
Protembia

Ptilophyllum pectcu ...

Pusiula

1 . 2

24,2

,26,2

20, 30
3

209

201

175

177
17

178
177

28, 47
45

12. 54
11
12

9, 12
13
52
20
21
38
21
21
21
21

13. 16
139
204

29

Quartzite Pre (’a7nbrian. of Toodyay district

88

Kaggatt, H.G.

Ramipora ambrosoides
Reed, F. R. C.

Refractories, sillimanite and andalusite
R ichthofen i idae
Rothpletz, A.

29

193

16. 26. 45
129

n

. 17. 26

General Index.

xciil.

l^owley. Pv. P.

Page

41

Salt Kango. India
Scacchinella

Schist, various mica, Toodyay district ...
Schmidt, H. ...

Schuchert, C.

Serpentire. Toodyay dif-tri(t
Serpentine, analysis ...

Silliinanite. ocHirrence in Toodyay district
Soils of Western Australia
Sowerby, .1. ...

SireptorhynchU’'< luiiuigvi
Sphenopterid fossils ...

Strophalo.'iiO' ...

Sfrophalosia rf. beecho.ri
Strophaloma clarkei ...

Strophalo-'sia comphctevs

Strophalosia etheridgei, ip. nf>v

Sirophalosia frayiUs ...

StrophaloMa (ferardi ...

Strophalosia cf. (jerardl
Strophalosia (jujas
Strophalosia horrescens
Strophalosia juke.si ...

Strophalosia kimberleyensis, s]>. nov.
Strophalosia lamellosa var. hvwblefov(/Ksi.s
Strophalosia multispinifera, sp. nov.
Strophalosia rarispimi
Strophalosia fenuispiva
Strophalosia sp. A. ...

Sirophalsiella
Slrophalosiitia
Strophalosiinae
SfrophoviP-va ...

Sutton, A. H.

S])inifex j)indan
Spirifer marvoni

7

12 ,

.. 9. 1(*
11
07

n

4. 10. 13. 29
128
128
08, 129
.. xxix.
30
22

.. 203

32, 37
41

41, 42, 45, 50
56

7. 41, 43, 52
.. 24, 39
, 45, 47
46
34

.. 34, 39
38, 47, 50
18, 41, 47, 50
38, 30

50
52

51, 52

51
40

35. 36, 40
9, 12, 53

9

... 1 I, 12

162
56

41, 4;

Taeniopleris elongata
Taeniopteris carruthersi
Taeniopieris spatvlata,

Taeniothaerus

Taeniothaerus snbqvadratvs ... ... ... ... ... ... 4, 7, 27

Talbot’s Cairn
Tayuan series
Teguliferina ...

Thinnfeldia

Thinnfeldia tabragare'nsis
Thomas, .1.

Tillighary, N.S.W

Tillyardewbia
Timor ...

Toodyay, geology and petrology of disti’ict of ...

Triodia pungens, Ecological succession observed during regeneration after burning
Triodia pungens^ Effect of burning on
Triodia ptingens, regeneration of ...

Tschernyschew, T. H.

. 201
. 203

. 203

27

28, 38
47
16
II

. 201
. 203

31
21
139

. 16, 26
83

149, 154
149
. 152

23

Cpper I^alaeozoic Hryozoa from Western Austi'alia ... ... ... ... ... ]H 7

Vegetation of Western Australia, Tlie

xl.

XCl V.

( Ienkrat. {ndk>^.

Waagen, L. ...

\y aagenoconclia
\y aagenoconcha imperfecta ...
Waagenoconcha vagans
Waltharrie Pool

Wandagee and Nooncanbah ISeries
Wandagee Station
Weller, S.

Western Australia, The \"egetatioi
Whitehonse. F. W. ...

\yinnemia

Permian) of
of

AVes1

ern Australia

Page

10, 12, 17, 18. 10. 2.'), 26, r>2

2o
26

41

Bryozoa from 165
31, 36, 43. 47. 54

42

xl.

Winning Station
Wyatkiiia
AVyndham Ga])
Wyndhamia

24
35. 36
18
5. 40

Yetar Spring ...

70

liy Authority: Uobkrt H. .\iiu.kh. (loveriinient Printer. Hertli.

*-

y'.-' yj%- i-.

mm

1

y