ASPECTS OF SCIENCE

IN

WESTERN AUSTRALIA

A volume to commemorate the 150th Anniversary of
the founding of Western Australia

Journal of the Royal Society of Western Australia, Vol. 62, Parts 1-4, 1979.

THE

ROYAL SOCIETY

OF

WESTERN AUSTRALIA

PATRON

Her Majesty the Queen

VICE-PATRON

His Excellency Air Chief Marshal Sir Wallace Kyle, G.C.B., K.C.V.O., C.B.E., D.S.O.,
D.F.C., K.St.J., Governor of Western Australia

COUNCIL 1978-1979

President ..

C. F. H. Jenkins, M.B.E., M.A., M.A.I.A.S.

Vice-Presidents M. J. Mulcahy, B.Sc. (For.), Ph.D.

J. R. de Laeter, B.Ed.(Hons.), B.Sc. (Hons.),
Ph.D., FJnst.P., F.A.I.P.

Past President

Joint Hon. Secretaries

Hon. Treasurer ..

Hon. Librarian .

Hon. Editor

A. J. McComb, M.Sc., Ph.D.

G. Perry, M.Sc.

M. W. Perry, B.Sc. (Agric.) (Hons.)
S. J. Curry, M.A.

A. Neumann, B.A.

A. E. Cockbain, B.Sc., Ph.D.

B. J. Beggs, B.Sc. (For.), Dip. For.

B. K. Bowen, B.Sc.

W. C. Ferguson, B.A. (Hons.)

J. K. Marshall, B.Sc. (Hons.), Ph.D.

L. J. Peet, B.Sc., Dip.Val., Dip.R.E.M., F.G.S., A.R.E.I.

P. E. Playford, B.Sc., Ph.D.

J. C. Taylor, B.Sc., Ph.D., A.R.C.S.

P. R. Wycherley, O.B.E., B.Sc., Ph.D., F.L.S.

ASPECTS OF SCIENCE

IN

WESTERN AUSTRALIA

1829

1979

A volume to commemorate the 150th Anniversary of
the founding of Western Australia

Journal of the Royal Society of Western Australia, Vol. 62, Parts 1-4, 1979.

ASPECTS OF SCIENCE IN WESTERN AUSTRALIA

Preface

When the early settlers landed in Western Australia in 1829 their first
priority was to wrest a living from the new and often hostile environment.
They had little inclination or opportunity to study the unique flora and fauna
which surrounded them and gave the bush creatures such misleading and
unimaginative names as native cat, native squirrel and native porcupine
and the commonest black and white bird was dubbed a magpie.

However, as more and more new plants and animals were found by explor-
ers and collectors popular interest was kindled and by 1891 the Premier Sir
John Forrest sponsored the formation of the Western Australian Natural
History Society.

This Society was short lived and became defunct in 1895, but again with
impetus from Sir John Forrest it was succeeded in 1897 by the Mueller Botan-
ical Society which was later expanded to embrace all forms of natural history
and laid the foundations for the Royal Society of Western Australia.

Since receiving its official Charter in December 1913 the Royal Society has
been very careful to comply with section two of its constitution —

“To promote and assist in the advancement of science.”

The 61 volumes of the Journal of the Royal Society of Western Australia so
far published cover a wide range of scientific disciplines and have earned for
the Society a very high reputation amongst scientists in many parts of the
world.

In presenting a special publication to commemorate the 150th Anniver-
sary of the Founding of Western Australia the Royal Society is proud of the
fact that many of its members have played an important part in the develop-
ment of the State and that some of the most important research findings first
appeared in its Journal.

The Society is grateful to the State Government for supporting its work
in past years and looks forward to the day when it will celebrate its own 150th
anniversary.

C. F. H. Jenkins,

President.

Journal of the Royal Society of Western Australia, Vol. 62, Parts 1-4, 1979, p. 3-11.

Aspects of Science in Western Australia 1829-1979.

A volume to commemorate the 150th Anniversary of the founding of Western Australia.

History of geology in Western Australia

by J. H. Lord

Geological Survey of Western Australia, 66 Adelaide Terrace, Perth, W.A. 6000

Abstract

Geology was of great interest to the early settlers because they realized that,
if their new colony was ever to develop rapidly, something in addition to agriculture
was required; hence the interest in locating economic mineral deposits. In the first
70 years geology was practised mainly by explorers and rarely by professional
geologists, which the colony could only occasionally afford to employ. This situation
changed with the first major mineral discovery — gold. Each major economic advance
in Western Australia has stemmed from new mineral discoveries. This has resulted
in the expansion of interest in and development of geology in government, academic
and industrial fields.

The beginning

In the development of Western Australia
during the past 150 years all of the periods of
major economic growth can be related to mineral
discoveries. Geology has therefore played a
major role in this development. Most of the
original settlers came with expectations of a
fertile land from which their own requirements
could be met by agriculture and animal hus-
bandry, but they soon realized that, apart from
relatively small areas such as along the Swan
and Greenough River valleys, the land was poor
and their methods yielded a bare subsistence.
Obviously something more was required to
attract new settlers and to provide earnings to
finance roads, schools, hospitals, harbours and
other essential public works.

Fortunately among the settlers were men of
vision who were prepared to look further afield,
and who became explorers; together with the
early surveyors, they ventured into the unknown
interior in search of better prospects. While
these persons were not trained geologists, they
were in many cases very good observers and
recorded the nature of the ground they traversed.
Also, because of the struggle for survival in these
early years, a few appreciated the advantages
which would accrue to the young colony if
economic minerals were discovered. As early as
1839 the Government offered a free grant of
1 036 hectares of land to the first person to dis-
cover a coal seam. Later a German naturalist,
Preiss, reported finding coal on the Murray River.
Specimens of this poor quality coal raised hopes,
and rumours of the mineral potential of the
colony were spread.

In September 1846, a prospectus to form a
company (The Western Australian Mining Com-
pany) with a capital of $40 000 was issued. This,
and the persistent rumours, generated the first
mineral boom for this State. To add to the

enthusiasm A. C. Gregory, the Assistant Surveyor
General, and his brothers, F. T. and H. C.
Gregory, reported the discovery of coal on the
Irwin River on 9 September 1846 and claimed
the free grant of land for the first discovery of
coal.

The company commenced drilling near the
Murray River and reached a depth of 12 m
encountering some coal of doubtful quality.
Through inexperience and shareholders who,
(like those of today) were looking for quick
returns, the company was soon in difficulties.
In order to assess the potential, the company
engaged Dr Ferdinand von Sommer, variously
described as “mining expert”, “eminent minera-
logist”, or “geologist”. His remuneration was
£6 per month with keep which amounted to
£12 19s 2d per month at Leeders Hotel and the
company horse, von Sommer arrived from
Adelaide on 15 February 1847, accompanied by
an experienced miner named Thomas.

The company was also interested in possible
lead or copper deposits between the Canning and
Serpentine Rivers, but the investigation for both
coal and metallic minerals was unsuccessful,
von Sommer’s employment with the Western
Australian Mining Company had lasted only a
few months. In mid- 1847 he was commissioned
by the Governor to accompany the Surveyor
General to examine the Gregory brothers’
coal find on the Irwin River. It appears that
von Sommer was the first professional geologist
to work in Western Australia and the first
employed by the Government. He travelled
extensively for those days, examining the
Victoria, Toodyay and York districts, the country
between York and Mount Barren and the area
from Doubtful Island Bay to Cape Naturaliste.

According to records this work was done in
1847 and 1848 and von Sommer wrote four
reports to the Colonial Secretary which are

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Journal of the Royal Society of Western Australia, Vol. 62. Parts 1-4, 1979.

recorded in exploration diaries of the Lands
Department. He contributed three papers to
the geological literature of the day and three
original drawings of his geological observa-
tions are preserved by the Geological Survey as
the first known geological maps of Western
Australia, von Sommer’s later activities are
unknown, and he left the colony in 1851.

The Gregory brothers, particularly A. C. and
F. T., both of whom were in the department of
the Surveyor General, carried out jointly and
separately many exploratory expeditions between
1846 and 1862, first in southern Western Austra-
lia, then in the North-West, and later across
northern Australia. They were keen observers
and their records frequently describe the geology
of the area over which they travelled. They
had a good knowledge of geology and contributed
papers to the Royal Geographical Society, of
which they were Fellows.

In 1848 F. T. Gregory discovered lead and
some copper in the bed of the Murchison River
and this occurrence, named Geraldine, became
the first lead mine in Western Australia. After
an extensive exploration trip through the North-
West, he reported “of mineral I was unable to
discover any traces, except iron”. A geological
map of the colony by the Gregorys was published
in London in 1860.

Due to the financial difficulties experienced
by the young colony, it was not until 1870 that
it was again possible to engage a geologist,
namely H. Y. Lyle Brown. It would appear that
he was given a two-year contract to report on
the geology of the colony, as then settled or
known. In his period of service Brown prepared
three maps and ten reports on the areas he
examined, which extended from the Murchison
southwards along the coast to Doubtful Island
Bay, and inland to the extent of the settled
area. A geological map of this area was pub-
lished on a scale of about 18 miles to an inch.
He later became the Government Geologist of
South Australia <1882-1912).

In April 1870, a clergyman/scholar the
Reverend Charles Grenfell Nicolay arrived in
Western Australia. He was a man of many
parts with the ability to antagonize some people
and please others. Nicolay, who was a Fellow
of the Royal Geographical Society, had been
associated with the King’s and Queen’s (now
defunct) Colleges in London, having been largely
responsible for founding the latter, where he
was Professor of Geography and Ancient History.
In Western Australia he apparently impressed
the Governors of the day, and, besides his church
duties, he became an adviser to the Government
(1881-1889) on technical and scientific matters.
His contributions to geology included an investi-
gation of a ship canal from the sea to Rocky
Bay on the Swan River to develop a major
harbour at Fremantle (probably the first engin-
eering geology done in this State), an investiga-
tion of the reports of coal in the bed of the
Fitzgerald River, a “Handbook of Western
Australia” (1881), the geology of the railway
route between Guildford and Clackline (1887)
and a circular on prospecting for gold (1886).
He also prepared a rock and mineral collection

together with some notes on the geology of
Western Australia for the Colonial and Indian
Exhibition of 1886, a contribution to a commis-
sion enquiring into the water supply of Perth
and Fremantle, and identifications of rocks and
minerals. In 1881 he was authorized by
Governor Robinson to establish at Fremantle a
collection of rocks and minerals of the colony.
Nicolay continued his many and varied interests
in the colony until his death in 1897 at the age
of 81.

After the departure of H. Y. Lyle Brown it
was nine years before the colony could afford
to employ a geologist again. E. Q. Hardman of
the Geological Survey of Ireland was appointed
in 1882 and stayed until 1885 when the colony
again could no longer afford the services of a
geologist. Hardman concentrated his efforts on
the Kimberley region and published two major
reports on his field work. These stimulated the
search for gold in this area, resulting in the
first find of gold in the colony by Hall and
party in 1885. The Government had offered a
reward of $20 000 to the discoverer of the first
goldfield to produce 10 000 ounces. While the
Kimberley find did not produce the required
amount of gold, the Government paid $1 000
reward each to Hardman and Hall. Hardman
also examined and reported on the geology of
the Bunbury and Blackwood River areas and on
the water supply of Perth and Fremantle.

Hardman made a major contribution to the
early geology of the State and returned to
Ireland with an assurance that he would become
Government Geologist when the Government
could afford to make a permanent appointment.
Unfortunately he died early in 1887, the year in
which it was decided to make such an appoint-
ment.

H. P. Woodward, who had been Assistant
Geologist to H. Y. Lyle Brown in South Austra-
lia, was appointed Government Geologist for
a term of five years and commenced duty in
January 1888. He had B. H. Woodward
appointed as assayer and curator of the
Geological Museum, while in 1894 S. Goczel was
appointed as Field Geologist and Mining
Engineer.

This was a very exciting period for Western
Australia with gold discoveries being made
progressively throughout the colony, commencing
in the Kimberley in 1885 and culminating with
Kalgoorlie in 1894. Woodward covered large
areas of Western Australia and produced short
reports on his observations for the Government.
His major work was probably the Mining Hand-
book issued in 1894 with a second edition in
1895. This volume, 216 pages with maps, must
have been extremely useful at that time because
as well as a description of the physiography and
geology, it contained a wealth of other informa-
tion including the known mineral occurrences
and resources, general information for prospec-
tors, and shipping, railway, postal and telegraph
information. Another accomplishment of
Woodward was the first coloured geological
map of the State published in 1894 by the
Government.

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Journal of the Royal Society of Western Australia, Vol. 62, Parts 1-4, 1979.

The colony began to appreciate the value of
competent geological advice through the efforts
of Brown, Hardman and Woodward, particularly
with the discovery of gold, which caused the
first major development. Woodward’s contract
was extended to the end of 1895 when he
resigned, as did his two assistants. Apparently
there were better-paid positions available else-
where in private enterprise. This resulted in
a firm decision being made to establish a
permanent Geological Survey in this State.

Until this period the knowledge and develop-
ment of geology had been confined to the
occasional Government Geologist employed by
the colony and to dedicated explorers or sur-
veyors, who had acquired a knowledge of geology
by reading and experience. From this time
geology was to be developed in Western Australia
by three groups of geologists: those associated
with Government Geological Surveys; the
teachers and research workers in educational
institutions; and consultants or geologists
employed by companies.

Government geological surveys

Andrew Gibb Maitland was appointed as
Government Geologist in charge of the first
permanent State Geological Survey. He started
work in November 1896 and established an office

Figure 1.— Andrew Gibb Maitland, the first Government
Geologist in charge of a permanent Geological Survey.

with a staff of nine, including three geologists,
a mineralogist, a topographic surveyor, a drafts-
man, a laboratory assistant, a cadet and a clerk/
accountant.

By 1912 the professional staff had increased
to ten; but later it decreased, until during the
1930s it averaged only three. It did not rise to
ten again until 1959. From 1961 onwards a
major expansion was approved and the number
of staff grew rapidly until at the end of 1978 the
establishment was 63 professional officers. The
Government Geologists (now titled Director) in
charge over this period were : Andrew Gibb
Maitland (1896-1926, Fig. 1); Torrington Blatch-
ford (1926-1934); Frank G. Forman (1934-1944);
H. A. (Matt) Ellis (1945-1961), and J. H. (Joe)
Lord (1961-present).

Until 1961 the geologists practised in all fields
of geology, with emphasis on geological mapping
and examination of mineral resources. It was
excellent general training for geologists, but
without opportunity to specialize in any particu-
lar field. After the expansion in 1961, geologists
were recruited for their particular experience or
training in specific fields of geology. The Survey
now has divisions covering most facets of geology,
namely hydrogeology (13 geologists), engineering
geology (4), mineral resources (9), regional
geology (9), sedimentary geology (9) and sections
in petrology (3), palaeontology (3), geophysics
(2), geochemistry (1), environmental geology (2)
and technical information (4).

Following the 1961 expansion of the Geological
Survey special emphasis has been placed on
regional geological mapping to ensure that the
geology of the whole State is mapped at the
useful 1:250 000 scale. The field work for the
last of the 163 maps should be completed in
this 150th Anniversary year. In this arid State
hydrogeology has become a major function of
the Survey in the search for groundwater for
town supplies, mining operations, and for farmers
and pastoralists. In the future more detailed
geological mapping is proposed for areas covering
mineralized belts.

Over the last 18 years, while efforts have been
mainly directed towards systematic geological
mapping, the Survey may have lost close
contact with the prospector and exploration
companies in the field, but this was necessary
if the mapping were to be done in a reasonable
time. To help restore this contact it is now
intended to open regional offices at several
major country centres.

The Commonwealth Government first became
involved in the geology of this State with the
passing of the Northern Australian Survey Act
of 1934. which gave authority to establish the
Aerial, Geological and Geophysical Survey of
Northern Australia (AGGSNA). The first meet-
ings of this organization were held in June 1934
and field parties started work in April 1935.
AGGSNA consisted of the Commonwealth
(Northern Territory), Queensland and Western
Australia, who agreed to start the three year
programme of work in each region north of 22 °S
latitude. The Commonwealth contributed
$150 000 and each State $75 000 to search for

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Journal of the Royal Society of Western Australia, Vol. 62, Parts 1-4, 1979.

metallic minerals, particularly gold. A geological
party was established in each State and two
geophysical parties which investigated problems
as and where required. The programme in
Western Australia extended beyond the planned
three years with field work being completed in
1938. The unit was disbanded in June 1939.

Some 64 reports on the Western Australian
work were published. The young geologists
involved in AGGSNA were later to become
leaders in the profession. Dr W. G. Woolnough
was the Commonwealth geological advisor for
the creation of AGGSNA and there is little
doubt that his enthusiasm was largely respon-
sible for the Commonwealth Government estab-
lishing the Bureau of Mineral Resources (BMR)
immediately after World War II

The BMR has been a major contributor to the
geology of this State, particularly prior to the
expansion of the Survey. Earlier the BMR
carried out all geological mapping in the sedi-
mentary basins while in more recent years it
has contributed to joint mapping parties. In
the field of geophysics the BMR has been
particularly active with regional aeromagnetic
and gravity surveys.

Educational institutions

As interest in geology heightened with the gold
discoveries of the 1890s, a demand arose for the
subject to be taught at the limited educational
facilities available at that time. In 1900 Perth
Technical School offered courses in geology and
mineralogy and these were probably the first
formal classes established. The lecturer was
P. C. Stockwell.

The School of Mines of Western Australia,
under the control of the Department of Mines,
was opened at Coolgardie in November 1902,
and then was moved to Kalgoorlie at the end
of 1903. Geology and mineralogy were amongst
the first courses offered. The School of Mines,
because of its locality, became a centre for
mining geology, although it was not until 1947
that a full associateship course was established.

The lecturer who probably deserves most
recognition is Dr C. O. G. Larcombe, who joined
the School in 1907 and remained there until
early 1934. He was responsible for planning the
Mineral Museum which opened on 7 December
1907, and is a feature of the School of Mines
today. Larcombe was particularly interested
in petrology and mineralogy, and he also acted
as petrologist to the Geological Survey of
Western Australia from 1922 to 1928. His best
known publication was “The Geology of Kalgoor-
lie with Special Reference to the Ore Body”
published in 1911 by the Australasian Institute
of Mining Engineers.

The School has always maintained an active
interest in the geology of the Kalgoorlie area.
Larcombe contributed many papers on this topic,
while Dr P. L. Stillwell, on loan from the Com-
monwealth Scientific and Industrial Research
(CSIR), used the School as a base for his study
in 1927 and 1928 of the “Geology and Ore

Deposits of the Boulder Belt, Kalgoorlie”
published as Geological Survey Bulletin 94. In
later years G. S. Compton and W. H. Cleverly
were in charge of the geology department.
When the Western Australian Institute of
Technology (WAIT) took control of the School
of Mines at Kalgoorlie in 1969 the Geology
Departments of the two establishments were
amalgamated, with the head of the Geology
Department being at WAIT near Perth.

WAIT was established in 1967 with an
associateship course in geology. This course,
which was upgraded to a degree course in 1975,
is designed to educate in the practical aspects
of geology, but there has been insufficient time
as yet for the graduates to make their mark.

When a university was first founded in this
State, limited finance restricted its scope to the
establishment of only seven chairs, exclusive of
Agriculture which was endowed. There was
much discussion and lobbying as to which
disciplines should be represented. The first six
decided upon were Mathematics and Physics,
Mining (now dropped) and Engineering, Biology,
History and Economics, English, Chemistry,
leaving the seventh chair to be either Classics,
Education or Geology. Voting was eventually
deadlocked between Classics and Geology, leaving
the Chancellor, Sir Winthrop Hackett, a well
known supporter of the Classics, with the casting
vote. To the Senate’s surprise the Chancellor
gave his vote to Geology because of the import-
ance of the mining industry to this State, a fact
which was realized in 1912 but is often over-
looked today.

By the time the University opened in 1913,
Walter George Woolnough (Fig. 2) had been
appointed the first Professor of Geology at the
University of Western Australia. He had studied
at the University of Sydney under the father
of geology in Australia, Professor Sir Edgeworth
David, and had gained the first D.Sc. awarded
by that University. He travelled extensively in
Australia and Fiji and had lectured at the
Universities of Sydney and Adelaide. As the
first geology professor in this State he was an
excellent choice, being a lucid lecturer and
devoted to his students to whom he gave a good
grounding in practical geology.

He resigned in 1919 and worked on economic
mineral deposits in all areas of Australia. In
1927 he was appointed Geological Adviser to the
Commonwealth Government, a position he was
reluctant to leave on reaching retirement age
in 1941, having laid the foundations for the
development of the Bureau of Mineral Resources.
He pioneered the use of aerial photography for
geological purposes in Australia, gave close
attention to the oil possibilities of the north
west of this State, in particular the Exmouth
Gulf area, and was responsible for the investiga-
tion of Yampi Sound iron ore deposits.

The Geology Department of the University
had been given a solid foundation by Woolnough,
but on his resignation it was decided, due to the
shortage of finance, to abolish the chair and to
appoint a lecturer-in-charge instead. This

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Journal of the Royal Society of Western Australia, Vol. 62, Parts 1-4, 1979.

Figure 2.— Walter George Woolnoueh, the first professor
of geology at the University of Western Australia.

decision allowed the estimated expenditure for
the geology department to be reduced from
$3 176 in 1919 to $1 080 in 1920.

Edward de Courcy Clarke was appointed to
the position of Lecturer but did not commence
duty until 1921. He was a graduate of Univer-
sity College, Auckland, and had joined the
Geological Survey of Western Australia in 1912
as a field geologist. He travelled extensively in
Western Australia and produced six bulletins
regarding his investigations on the goldfields,
and had made a reconnaissance survey with H.
W. B. Talbot of the country between Laverton
and the South Australian border. He enjoyed
field work and practical geology, spending 356
days in the field around Meekatharra in 1914.

As a lecturer his enthusiasm for practical
geology was always evident and he was a worthy
successor to Woolnough. In 1930 the chair of
geology was re-established; Clarke was appointed
and occupied it until his retirement in 1948. He
was a dedicated man with simple tastes and an
ability to encourage and inspire his students,
whose interests he placed above all others. He
consolidated the Geology Department and estab-
lished its reputation. His main interest was
Precambrian geology, although in the latter

part of his career he was co-author with his
staff of many papers on the geology of younger
rocks.

Clarke was succeeded by one of his outstanding
students, Rex T. Prider, who carried on the
traditions established by Clarke until he retired
in 1975. Peter G. Harris, another New Zealand
graduate, now occupies the chair.

During the past 30 years the Geology Depart-
ment has gradually expanded to meet the
demand for more geologists and research. One
might expect that this Department of the major
University in Western Australia would, because
of the huge mineral wealth in the State, have
developed into a major centre for research into
the Precambrian rocks and mineral deposits.
This has not eventuated although such centres
have now developed in other Australian Univer-
sities.

Consulting and company geology

The discovery of the Golden Mile near Kal-
goorlie in 1894 created world-wide interest in
Western Australia, resulting in visits by overseas
consulting geologists and mining engineers, as
well as government and academic geologists.
Many of these persons wrote of their observa-
tions and opinions. Amongst those who
published papers within the first ten years of the
discovery were: G. J. Bancroft, H. Y. L. Brown,
Y. F. Bulman, A. F. Calvert, S. W. Tard, A.
Carnot, A. G. Charlton, C. Chewings, H. B.
Corbin, William Frenchville, E. Halse, H. G.
Holroyd, Herbert C. Hoover, J. F. Kemp, P. M.
Krause, P. Krusch, Baron Sloet van Oldriuten-
burgh, E. F. Pitman, T. A. Rickard, Bergrath
Schmeisner and Brenton-Symons. After the
initial flurry, from about 1910 onwards more
detailed geological examinations were made by
consultants, who stayed longer. Many of their
reports were no doubt confidential to the com-
panies concerned. Amongst these were Dr
Malcolm McLaren (representing the London
Mining House, Gold Fields), Dr J. A. Thomson,
R. Lockhart Jack and T. Esdaile.

Due to World War I and the depressed con-
ditions for gold in the aftermath, little work is
recorded by private consultants or companies in
this period. In 1928 Malcolm McLaren returned
to look at the Lake View and Star Mine at
Kalgoorlie and the Moonlight-Wiluna Gold mine
at Wiluna. Probably through him a Geological
Section was established at the Lake View and
Star Mine in 1929 with a young geologist from
Canada, R. F. Playter, in charge, assisted by
Harold Jensen. This was probably the first
company geological office established in this
State.

In 1933 Western Mining Corporation (WMC)
was formed by William Sydney Robinson, who
was a strong supporter and instigator of a
scientific approach to exploration. This has
certainly been proved profitable when one looks
at WMC’s existing mines and prospects (Yeel-
irrie, Roxby Downs and Benambra) . A
Geological Department was established with J.
D. Campbell, H. W. B. Talbot, A. Blatchford and
K. J. Finucane as the first geologists.

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Journal of the Royal Society of Western Australia, Vol. 62, Parts 1-4, 1979.

Robinson then engaged D. McLaughlin,
Professor of Economic Geology at Harvard, as
consultant to the company to introduce modern
methods of exploration, which he had developed,
into Western Australia. McLaughlin assembled
a large group of geologists who had trained
under him at Harvard. H. E. McKinstry, G. K.
Gustafson and F. Miller were consultants for
the company from 1934 until 1936, while H. J. C.
Connolly was appointed chief geologist and
Haddon F. King, Frank A. Moss and R. F. Thyer
joined as permanent staff members. This prob-
ably still remains the most talented group of
geologists ever assembled for mineral exploration
in Australia and WMC retained more geologists
than any other company operating elsewhere
in Australia at that time.

WMC was the first organization to apply
aerial photography to mineral exploration in
Australia. This technique had been suggested
by South African interests in the company, and
the photographic section was under the control
of Wing Commander Laws, a former Royal
Flying Corps pilot. Some of the experience and
techniques developed were used in World War II.
The group carried out factual surface mapping
and interpretation. Geophysical methods were
considered and ground magnetic surveys were
done.

In addition, the group examined existing and
former mines for extensions or repetitions.
These included the southern extension of the
Golden Mile, Norseman, Great Fingall (Day
Dawn) and the Cosmopolitan (Kookynie).
Through pursuing this policy with an active,
progressive geological team, WMC has become
one of Australia’s major mining companies, and
with such finds as mentioned earlier, it has
proved that geologists as well as prospectors do
find ore bodies.

After World War II, other companies such as
BHP, Rio Tinto and Zinc Corporation, began to
employ exploration geologists in this State and
in the oil and uranium booms of the early 1950s,
companies and consultants established geological
departments. In the mineral boom of the 1960s
and early 1970s geological sections became
essential for all active companies, some of the
larger companies having 60 to 80 geologists
engaged at the peak of the boom.

While a number of geological consultants
visited this State for varying periods, it is
believed that K. J. Finucane (Fig. 3), a Western
Australian graduate with wide experience, was
the first to establish himself as a permanent
resident consultant in 1939. He is still practising
some 40 years later. In the early 1950s Frank
G. Forman, a former Government Geologist,
established a consultancy; he was followed by
E. P. Utting and S. A. Tomich later in the
decade.

The mineral boom of the 1960s saw many
local and overseas individual consultants and
consulting firms establish themselves in Western
Australia. With the passing of the boom in 1972
some moved on, some have turned to contract
work or positions with companies, while about
30 continue to practise.

Figure 3.— Kevin J. Finucane, the first resident consult-
ant in Western Australia.

Specialized branches of geology

With the rapid development of geology in the
last 2 or 3 decades many new branches or
associations with geology have developed parti-
cularly in the search for minerals. These include
geophysics, geochemistry, geobotany, palaeo-
magnetism, geochronology and others, which
will eventually require a history of their own.
Only the original major branches of geology
are considered below.

Petrology

Petrology has a longer history in this State
than any other branch of geology. This resulted
from the great interest developed here and
abroad with the discovery of gold at Kalgoorlie,
and the necessity to unravel the petrological and
structural history, a problem which is still
receiving attention.

Many geologists visiting the new goldfields
found great interest in the petrology and miner-
alogy of the telluride ores and their host rocks
which led to the publication of numerous short
papers while no doubt many others were held
as confidential in company reports.

The first published descriptive petrology was
by G. W. Carr of the Geological Survey of New
South Wales on a suite of rocks from Kalgoorlie.
However E. S. Simpson, although strictly a
mineralogist and analyst, had joined the
Geological Survey in 1897 and was contributing
to petrological knowledge.

8

Journal of the Royal Society of Western Australia, Vol. 62, Parts 1-4, 1979.

About 1910 microscopic petrology flourished
with Dr J. A. Thomson, who accompanied Dr
Malcolm McLaren on a lengthy consulting visit,
contributing papers. Dr C. O. G. Larcombe, at
the School of Mines, studied the rocks of the
Golden Mile, and R. A. Farquharson joined the
Survey as petrologist in 1911. After Farquhar-
son’s resignation in 1922, Larcombe acted as
petrologist for the Geological Survey until 1928.
The University was the next to develop petrology
with the appointment of Rex T. Prider as an
assistant lecturer in 1934. Later he went on to
Cambridge to study for his Ph.D. and returned
to the University, where he made many major
contributions to the petrology of this State. The
University continues to be well known for its
petrological research.

Sedimentary petrology was studied at the
University of Western Australia with the
appointment of Dr Dorothy Carroll in 1941 as
a Research Fellow. After her departure in 1943,
the University did not appoint a replacement
until Dr J. J. E. Glover joined as a Lecturer in
1955 and he has continued to study and teach
this branch of petrology.

After Farquharson’s resignation the Geological
Survey did not appoint another petrologist until
Dr A. F. Trendall joined the staff in 1962. The
Survey’s Petrology Section has now been
expanded to three petrologists. Over the last 10
years joint petrological and geochronological
studies with John de Laeter of WAIT have been
carried out and have made a major contribution
to this work in Western Australia (see de Laeter
and Trendall, this volume).

Palaeontology

This branch of geology in Western Australia
illustrates the development of specialization in
the science. In the last century there were
very few palaeontologists, but generally they
examined most groups of fossils. Today, in
contrast, palaeontology is a highly specialized
profession, with some workers spending their
lifetime studying a few genera of one particular
order. This applies particularly to the study of
foraminifers, due to their economic value in oil
exploration.

In Western Australia the early palaeontology
was of a very general nature and studied mainly
by overseas workers to whom suites of specimens
were sent. F. T. Gregory, over the period 1846-
1861, collected fossils particularly from the Irwin
and Gascoyne Rivers and Geraldton areas. The
Gregory brothers made the first fossil determin-
ations and showed the results on their geological
map of 1847. Other collections of fossils were
sent to England, and studied by Charles Moore,
Reverend W. B. Clarke and W. H. Huddleston
and reported upon in the Quarterly Journal of
the Geological Society of London.

In the latter part of the 19th century E. T.
Hardman and H. P. Woodward devoted some of
their time as Government Geologists to the study
of fossils and encouraged other specialists, such
as Arthur H. Foord (brachiopoda and mollusca),
R. Kidston (plant remains), H. Alleyne Nichol-
son (stromatoporoids) and George J. Hinde
(corals and bryozoa) to become involved.

With the establishment of a permanent Geo-
logical Survey, R. Etheridge Jr., Curator and later
Director of The Australian Museum, Sydney,
became the honorary palaeontologist and
together with others, including F. Chapman,
palaeontologist, National Museum, Melbourne,
and W. Howchin, palaeontologist. University of
Adelaide, had numerous papers published in the
Geological Survey’s bulletins.

In 1908 Ludwig Glauert was engaged by the
Survey as a palaeontologist, apparently the first
employed full-time in this State. He prepared
a catalogue of Western Australian fossils and
appeared to have stimulated more work by
outside specialists including E. A. Newell Arber
and Rex W. Bretnall. Unfortunately Glauert
resigned within two years to join the Western
Australian Museum, where he continued for
many years producing palaeontological papers
published mainly in the Journal of the Royal
Society of Western Australia.

The Geological Survey did not engage another
palaeontologist until 1962; the staff of the
palaeontology section has now increased to
three. The palaeontologists provide general
palaeontological assistance to the geological
staff; palynology is particularly useful for strati-
graphic control of the extensive groundwater
exploratory drilling programmes carried out by
the Survey.

The University of Western Australia did not
specialize in palaeontology until Dr Curt
Teichert was appointed a Research Fellow in
1937. During his eight years at the University,
Teichert was responsible for considerable
research in palaeontology and stratigraphy and
his papers drew world-wide attention to this
State and in particular to its oil potential. After
Teichert .eft, the University continued to have
palaeontologists on the staff, while palynology
was added later to its area of research and
teaching. Dr B. E. Balme, who had done an
Honours thesis on palynology of the Collie coals
in 1948, returned to the University some ten
years later to become a Lecturer (later Reader),
specializing in palynology.

In recent years the Western Australian
Museum has studied vertebrate and Quaternary
palaeontology.

Mineralogy

When the Geological Survey was formed
provision was made for a mineralogist and
assayer. In 1897 Edward S. Simpson, graduate
in mining and metallurgy from the University
of Sydney, was appointed to the position. He
was destined to stay with the Government for
42 years and in this time made a major and
oustanding contribution to his field.

As mineralogist and assayer he acted as
chemist to the Department of Mines. In addition
to handling work of a routine nature, he became
involved in the chemistry and petrology, as well
as the mineralogy, of areas being studied by
the Survey. In particular he unravelled the
rock types of the Golden Mile at Kalgoorlie.
In 1919 he was awarded a D.Sc. from the
University of Western Australia for hi s thesis

9

Journal of the Royal Society of Western Australia, Vol. 62, Parts 1-4, 1979.

“The Minerals of Western Australia”. Through-
out his life he continued to study and accumulate
information on the minerals of this State.

In 1922 he was made Government Mineralogist
and analyst in charge of what is now known
as the Government Chemical Laboratories, which
conducts a wide variety of analytical work for
numerous Government Departments. Despite
these added responsibilities, he continued to
promote and foster mineralogy, the study of
which he retained as a major operation in the
laboratories.

He published over 100 scientific papers, pro-
vided an immense amount of data for Survey
publications and was working on his opus
magnum summarising the mineralogical know-
ledge of the State when he died suddenly in
August 1939, only a few months prior to retire-
ment. His work was completed by colleagues
and “Minerals of Western Australia” was even-
tually published in three volumes by the Govern-
ment over the period 1948 to 1952. This has
proved to be an invaluable reference book on
the minerals of the State and their occurrence,
to assist students, prospectors and exploration
geologists.

Since the passing of Simpson and his immedi-
ate successors the study of mineralogy has
declined. The Government Chemical Labora-
tories have now grown to a size where they
could be decentralized to the Departments for
which they work.

As mineralogy forms an integral part of
petrology, geochemistry and geochronology, it
continues to be studied at the University of
Western Australia but there has been no strong
specialization in the subject.

In 1963 the Commonwealth Scientific and
Industrial Research Organization (CSIRO)
opened a branch of their Division of Mineralogy
in Perth. A very strong and active research
section on the problems of mineralogy and
mineralization in this State has been developed
and is contributing to our development.

Geological museum

Rev. C. G. Nicolay, started the first collection
of rocks, minerals and fossils at Fremantle in
1881. Nicolay was apparently very keen and,
established a good geological collection represen-
tative of the colony. To the best of his ability
he tried to identify and report on the rocks and
minerals that he collected or acquired. He did
receive criticism from one Government Geologist
of the day, E. T. Hardman, who apparently
considered him to be incompetent in his identi-
fications.

The collection assembled by Nicolay was
housed at Fremantle and known as the Geo-
logical Museum. In 1889 it was transferred to
Perth, to form the original collection of the
Western Australian Museum.

Several collections of minerals including
excellent specimens of gold and tellurides from
the Kalgoorlie area were sent to exhibitions in
England and Europe, but were never returned.

When Maitland formed a permanent Survey
in 1896, he and his staff built up a collection
of rocks and minerals. This collection far out-
grew the Museum collection, and in 1908 the
Government decided that the two collections,
both at that time housed close together, should
be amalgamated under the direction and custody
of the Government Geologist, but displayed in
a gallery attached to the Museum. In the
amalgamated collection there were about 10 000
registered specimens of which only about 15 per
cent had been derived from the Museum’s
collection.

The collection continued to grow until 1966
when the Museum required the gallery for other
purposes and the Survey was required to remove
the collection.

It was placed in store at the Survey’s core
and cuttings library until 1971 when Mineral
House was completed with limited space for a
Museum. Now nearly 1 000 specimens of Western
Australian rocks, minerals and fossils are
displayed in the Survey’s Museum, while the
remainder are still in store. The catalogue of
the whole collection is now being placed on
computer and retrieval of any particular speci-
men is relatively easy if required.

The Western Australian Museum has concen-
trated on a collection of vertebrate and Quatern-
ary fossils. It also has a collection of meteorites
which attracts world-wide recognition but no
collection of rocks or minerals.

Conclusion

This brief review of the development of
geology in the State shows that the profession
has gained greater strength in the last 20 years
than in the previous 130 years. This is the
result of the extensive mineral development of
recent times and the general acceptance that
geology is necessary to locate, to explore, and to
develop mineral deposits.

The growth of the profession in this State
appears at first glance to be very satisfactory,
until comparison is made with that of other
States. In the Western Australian Geological
Survey there is only one geologist per 40 000 km 2 ,
which is the lowest density of all the Australian
States. Queensland has one geologist per
24 000 km 2 , while Tasmania has one per 2 100
km 2 . Thus it is apparent that this situation can
and should be improved as soon as finance can
be made available. There is need for expansion
of teaching, both at secondary and tertiary
level, and research into many aspects of geology.
This State should become a centre for study of
both Precambrian geology (metallic minerals)
and sedimentary geology (oil and gas) by the
establishment of research centres, particularly
in association with the tertiary institutes.

As this State, and Australia generally, depends
largely upon minerals, it is the duty of the State
to arrange the financing of these proposals. If
this is done, our economic future will be assured
and at Western Australia’s bicentennial anniver-
sary, the State will be known as a centre of
geological research and geological activity, as

10

Journal of the Royal Society of Western Australia, Vol. 62, Parts 1-4, 1979.

well as having developed into the major mineral
production and processing region in Australia.

Acknowledgements. — I wish to acknowledge useful
discussion and information supplied by many people
and in particular, the following: J. D. Campbell, R. R.
Connolly, K. J. Finucane, F. G. Forman, J. J. E. Glover,
R. T. Prider and D. R. Reid.

References

Numerous references have been consulted but,
in consideration of space only the major ones
are listed. However from those given, all others
may be located if required. For instance, I. R.
Williams has assembled nearly 1 000 references
on the Eastern Goldfields of which many of the
earlier ones have been consulted but are not
quoted here.

Alexander, F. (1963). — Campus at Crawley. University of
West. Australia Press.

Battye, J. S. (1924). — Western Australia. Oxford Univer-
sity Press.

Geological Survey of Western Australia (1974). — Geology
of Western Australia: West. Australia Geol.
Surv., Mem. 2.

Glauert, L. (1910). — Palaeontological contributions to the
Geology of Western Australia. A list of
Western Australian Fossils. West. Australia
Geol. Surv. Bull. 36: 71-106.

Glauert, L. (1926). — Palaeontological contributions to the
Geology of Western Australia. A list of
Western Australian Fossils: West. Australia
Geol. Surv. Bull. 88: 36-71.

Gregory, A. C., and Gregory, F. T. (1884). — Journals of
Australian Explorations. Brisbane.

Hoover, Herbert C. (1899). — The superficial alteration of
Western Australian ore deposits: Trans. Am.
Inst. Min. Eng. 28: 758-765.

Johns, R. K. (Ed.) (1976). — History and role of Govern-
ment Geological Surveys in Australia: Govt
Printer, South Australia.

Maitland, A. Gibb (1898). — Bibliography of the geology
of Western Australia: West. Australia Geol.
Survey Bull. 1.

Maitland, A. Gibb (1911). — The origin, history and work
of the Geological Survey of Western Aus-
tralia: West. Australia Geol. Serv. Ann. Rept
1910 7-13.

Playford, P. E., and Pridmore, I. (1959). — The Reverend
C. G. Nicolay, Roy. West. Australia Hist.
Soc. J. & P. 7: 29-33.

Quilty, P. G. (1975). — An Annotated Bibliography of the
Palaeontology of Western Australia 1814-1974.
West. Australia Geol. Surv. Rept 3.

Richard, T. A. (1899). — The Alluvial Deposits of Western
Australia. Trans. Am. Inst, of Min. Eng. 28:
490-537.

Simpson, E. S. (1948). — Minerals of Western Australia.
Govt Printer, Perth.

Thomson, J. A. (1913). — On the Petrology of the Kal-
goorlie Goldfield (Western Australia). Quart.
J. Geol. Soc. 69: 621-677.

von Sommer, F. (1849). — A Sketch of the Geological
Formation and Physical Structure of Western
Australia: Quart. J. Geol. Soc. 5: 51-53.

Williams, I. R. (1974). — A review of the geological inves-
tigations of the eastern goldfields province,
including bibliography to December 1972.
West. Australia Geol. Surv. Rec. 1973/11.

Woodward, H. P. (1891). — Annual General Report for
the Year 1890. Govt Printer, Perth.

11

Journal of the Royal Society of Western Australia, Vol. 62, Parts 1-4, 1979, p. 13-20.

Aspects of Science in Western Australia 1829-1979.

A volume to commemorate the 150th Anniversary of the founding of Western Australia.

Stromatolite research in Western Australia

by Phillip E. Playford

Geological Survey of Western Australia, 66 Adelaide Terrace, Perth, W.A. 6000

Abstract

Research on Western Australian stromatolites, both modern and ancient, has
played an important role in the environmental and biostratigraphic interpretation
of these enigmatic organosedimentary structures. This research has been concentrated
on modern stromatolites from Hamelin Pool, Devonian stromatolites from the
Canning Basin, and Proterozoic stromatolites from several areas of the State.

The first research on the Hamelin Pool stromatolites indicated that they are
restricted to the intertidal zone, their morphology is controlled by environmental
rather than biological factors, and they are formed solely by the trapping and
binding of sedimentary particles by algal filaments. Using uniformitarian principles
these conclusions were widely applied to ancient stromatolites, and as a corollary,
many palaeontologists and stratigraphers concluded that stromatolites can be of no
value for biostratigraphic correlation. However, each of these conclusions has
subsequently been disproved or substantially modified.

Recent work at Hamelin Pool has now shown that living stromatolites there
are not restricted to the intertidal zone as had previously been supposed; they grow
to depths of at least 3.5 m. Indeed the subtidal occurrences are more widespread
than those in the intertidal zone. Moreover, although the external morphology of
stromatolites at Hamelin Pool is governed by environmental factors, their internal
fabric is biologically controlled.

Stromatolites in the Devonian reef complexes of the Canning Basin grew
through a wide depth range, from the intertidal zone to depths which probably
exceeded 100 m. These stromatolites were not formed solely by enmeshing of
clastic particles by algal filaments; biochemical precipitation of carbonate by algae
and/or bacteria was also important, and was the only growth mechanism in some
forms. The shallow- and deep-water stromatolites in these reef complexes have
certain distinguishing morphological features which may also characterize other
ancient stromatolites. The most distinctive of these is the presence of fenestral
fabrics in shallow- water forms (also characteristic of Hamelin Pool stromatolites)
and their absence in deep-water forms.

Western Australian Proterozoic stromatolites have been shown to have value for
inter-regional correlations within Australia. However, some significant anomalies
have been found in attempting to correlate with the standard stromatolite sequence
in the Proterozoic of the Soviet Union, and there remains considerable doubt as to
the extent to which stromatolites can be used for inter-continental correlation.

Introduction

Interest in the study of stromatolites has
increased greatly among palaeontologists, sedi-
mentologists, and biologists over the past 20
years, primarily because stromatolites are the
principal macrofossils known from the Precam-
brian, and are potentially useful for biostrati-
graphic correlation and environmental interpre-
tation.

However, there has been considerable
controversy on the validity of correlations and
environmental interpretations using stromatolites.
Some authorities have maintained that the
morphological characteristics of stromatolites
are environmentally rather than biologically

controlled, and that consequently stromatolite
taxa are meaningless in biological terms, and
biostratigraphic correlations based on them can
have no validity. These workers commonly
favour a stromatolite classification based on
descriptive geometric formulae. Others believe
that evolutionary changes must have occurred
in the stromatolite-building algae, thereby
resulting in recognizable changes in the stroma-
tolites themselves through geological time. They
claim that although environmental factors are
often important in controlling the gross external
morphology of stromatolites, their internal fabric
is largely biologically controlled. These authori-
ties use a binomial nomenclature of “group”
and “form” in classifying stromatolites. They

Journal of the Royal Society of Western Australia, Vol. 62, Parts 1-4, 1979.

acknowledge that the terms “species” and
“genus” should not be applied to stromatolites,
as they have commonly been built by assemblages
of algae and/or bacteria. Many of these workers
believe that stromatolites are of considerable
value for inter-regional and perhaps inter-
continental correlations in the Precambrian. A
recent comprehensive review of the status of
world stromatolite research is contained in the
volume edited by Walter (1976).

With regard to the environmental interpreta-
tion of stromatolites, some researchers have
claimed that marine stromatolites are restricted
to narrow environmental limits, being virtually
confined to the intertidal zone. Others believe
that stromatolites have grown through a very
wide range of environments, from the supratidal
zone to abyssal depths of the oceans.

Research on Western Australian stromatolites
has played an important role in the history of
debate on these critical aspects of stromatolite
studies, and it is to these aspects that I will
primarily direct this paper.

The term stromatolite as used in this paper is
defined as an organosedimentary structure with
vertical relief above the substrate produced by
sediment trapping and/or precipitation resulting
from the growth of micro-organisms, principally
blue-green algae (slightly modified after
Awramik 1977).

Historical review

The first description of stromatolites in
Western Australia was by Clarke and Teichert
(1946) dealing with wrinkled and bulbous
structures in algal mats covering the floor of
Lake Cowan, a dry salt lake near Norseman.
Although Clarke and Teichert did not use the
term stromatolite in describing these structures,
they would be referred to as such by modern
workers.

Fairbridge (1950) was the first to publish a
description of ancient stromatolites from this
State. He described Proterozoic silicified forms,
which he referred to the genus Collenia, from
what is now known as the Coomberdale Chert,
in the area north of Moora. Logan and Chase
(1961) subsequently described Collenia undosa,
C. columnaris, and Cryptozoon frequens from
this formation.

Logan (1961) published a paper dealing with
the modern stromatolites of Hamelin Pool, an
arm of Shark Bay; this was the first research
on Western Australian stromatolites to receive
international attention. The Hamelin Pool

stromatolites are now recognized as being the
best examples of living stromatolites known
from modern seas, and the conclusions reached
by Logan had a profound impact on stromatolite
research for many years. Several subsequent
papers have described these remarkable forms,
including those by Logan et al. (1974), Hoffman
(1976), and Playford and Cockbain (1976).

An attempt to use stromatolites for corre-
lation of Proterozoic stromatolite-bearing

sequences in Western Australia was published
by Edgell (1964). He concluded, contrary to

prevailing opinion in Australia at the time, that
biostratigraphic correlations could be made using
stromatolites, at least on an inter-regional level.

A varied assemblage of stromatolites and a
series of large stromatolite bioherms have been
described from the Devonian reef complexes of
the Canning Basin by Playford and Cockbain
(1969) and Playford et al. (1976). These are
among the best-preserved stromatolites known
from middle Palaeozoic rocks in the world.
However, their unique importance lies in the
fact that it has been possible to demonstrate
the original water- depth relationships of the
stromatolites, and to formulate criteria for
distinguishing deep-water and shallow-water
forms.

Modern studies of Proterozoic stromatolites in
Western Australia began with the monograph by
Walter (1972), in which he described forms from
the Hamersley, Bangemall, and Ord Basins in
Western Australia, and from other Australian
basins. Walter applied methods of stromatolite
study developed in Russia, where stromatolites
have been studied more intensely than in other
countries and have been widely used for inter-
regional Precambrian correlations.

The most recent work on Proterozoic stromato-
lites in Western Australia is by Preiss (1976),
describing forms from the Nabberu and Officer
Basins. This work is important principally
because of the doubt that it casts on the validity
of inter-continental correlations in the Proter-
ozoic based on stromatolite groups.

Modern stromatolites at Hamelin Pool

Introduction

Hamelin Pool is a marine barred basin con-
sisting of a broad central area 5 to 10 m deep,
surrounded by a sublittoral platform up to 5 km
wide, backed by intertidal-supratidal flats. The
basin is barred to the north by the Faure Sill,
a sand and seagrass bank cut by a number of
prominent tidal-exchange channels. The high
evaporation and low precipitation in the area,
combined with the restricted inflow of normal
oceanic water because of the Faure Sill, has
caused the waters of Hamelin Pool to become
hypersaline, with salinities ranging from 55°/oo
to 70°/oo throughout the year.

Stromatolites were first recognized around the
shores of Hamelin Pool by geologists of West
Australian Petroleum Pty Ltd in 1954-55, and
the first published description of them was by
Logan (1961). This paper and the subsequent
paper by Logan et al. (1964) stimulated interest
in the study of stromatolites around the world,
and their conclusions greatly influenced the
interpretation of ancient stromatolites. As a
result it became widely accepted that stromato-
lites are essentially confined to the intertidal
zone, that their morphology is environmentally
rather than biologically controlled, and that they
have formed solely by the trapping and binding
of particulate sediment by algal filaments without
any biochemical precipitation. Moreover, because
of the supposed lack of biological control on
stromatolite morphology, it was generally

14

Journal of the Royal Society of Western Australia, Vol. 62, Parts 1-4, 1979.

believed that biostratigraphic correlations based
on stromatolite taxa could have no validity.
However, each of these conclusions, founded
largely on the early work at Hamelin Pool, has
since been disproved or substantially modified.

Distribution

Living stromatolites extend discontinuously
around the shores of Hamelin Pool from the
intertidal zone to depths of at least 3.5 m on
the sublittoral shelf. Logan (1961) had claimed
that the Hamelin Pool stromatolites are confined
to the intertidal zone, and that as a result the
maximum height of individual columns cannot
exceed the mean tidal range (about 0.6 m). This
conclusion was confirmed by Logan et al. (1964)
and was extended to embrace stromatolites in
the ancient record. However, subsequent work
(first reported by Playford 1973) has shown that
subtidal stromatolites are widespread at Hamelin
Pool.

Logan et al. (1974) reported that large subtidal
stromatolites occur at a few localities in Hamelin
Pool, extending to depths of about 2 m, but
concluded that these forms could be “relict”
structures which grew originally in the intertidal
zone at a time when sea level was lower than
it is today. Hoffman (1976), in a summary of
this paper, also recorded subtidal stromatolites
to depths of 2 m, but he did not imply that they
were relics of an earlier low sea level.

Playford and Cockbain (1976) observed that
contrary to previous reports, living stromatolites
and flat algal mats are widespread over the
sublittoral platform fringing Hamelin Pool, and
that the stromatolites extend to depths of at
least 3.5 m. Furthermore, subsequent observa-
tions utilizing low-level colour air photos show
that subtidal stromatolites and algal mats at
Hamelin Pool are more extensive than the
intertidal forms.

In many areas of Hamelin Pool living stroma-
tolites are backed by dead forms in the supratidal
zone, and these reach 0.5 to 1 m above mean sea
level, commonly forming a series of stepped
terraces (Playford and Cockbain 1976). These
dead stromatolites are in varying stages of
disintegration, and they must have emerged
in relatively recent times (perhaps only several
hundred years ago). It is uncertain whether
this emergence has resulted from tectonic uplift
or eustatic falls in sea level, but there is increas-
ing evidence for Holocene tectonism in this part
of the Carnarvon Basin, and tectonic emergence
of the dead stromatolites is therefore likely.

Morphology and biology

Logan (1961) emphasized environmental con-
trols on stromatolite morphology at Hamelin
Pool, but he did not distinguish the various types
of stromatolite-building algal mats and the
internal fabrics resulting from those mats. It
was later shown by Logan et al. (1974) and
Hoffman (1976) that there are three types of
algal mat that build stromatolites at Hamelin
Pool: pustular mat in the middle to upper inter-
tidal zone, smooth mat in the lower intertidal

zone, and colloform mat in the lower intertidal
to subtidal zone. Each of these mats is built by
a characteristic association of algae; pustular
mat is dominated by Entophysalis major, smooth
mat by Schizothrix helva, and colloform mat by
Microcoleus tennerimus. These authors also
showed that the internal fabrics of stromatolites
resulting from the three mat types are different.

These observations on mat distribution and on
the relationships between mat types and internal
fabrics were generally confirmed by Playford
and Cockbain (1976). However, more recent
observations show that pustular mat extends
from the upper intertidal zone into the
shallowest subtidal zone, smooth mat is charac-
teristic of the middle to lower intertidal and
shallow subtidal zones (extending deeper than
pustular mat), and colloform mat is restricted
to subtidal environments, reaching depths of at
least 3.5 m. Thus there is a generalized zonation
of pustular mat dominating the shallowest
environments, followed successively moving into
deeper water by smooth mat and colloform mat.
Pustular mat results in massive or crudely
layered, irregular, coarse fenestral fabrics,
smooth mat in laminated fine fenestral fabrics,
and colloform mat in weakly laminated, coarse
fenestral fabrics.

The most characteristic feature of the Hamelin
Pool stromatolites is the occurrence of fenestral
fabrics, the largest fenestrae being found in
stromatolites formed by pustular mat, and the
smallest in those formed by smooth mat. The
best-developed lamination is formed by smooth
mat, while the poorest results from pustular mat.

Monty (1976) and Golubic (1976) give further
details of some aspects of the microstructure
and internal fabrics of Hamelin Pool stromato-
lites and of their biological characteristics.

Logan (1961) concluded that discrete stroma-
tolites occur characteristically around headlands,
whereas flat algal mats occur in bays, and this
was generally confirmed by Logan et al. (1974)
and Hoffman (1976). They observed that

stromatolites at Hamelin Pool are commonly
elongate in the direction of wave translation,
approximately normal to the shoreline, and
deduced that such elongation in ancient stroma-
tolites is likely to have a similar origin. They
also reported that some stromatolites “lean”
seaward in the direction of wave translation.
Hoffman (1976) noted further that stromatolites
in some areas of Hamelin Pool occur in belts
parallel to the shorelines, but gave no explana-
tion for this alignment.

Playford and Cockbain (1976) found that
although intertidal stromatolites in some areas
are concentrated around headlands, in others
they are extensively developed in bays. They
also found that subtidal stromatolites are wide-
spread in front of both headlands and bays. It
now seems that the main requirement for the
extensive development of stromatolites at
Hamelin Pool is that there be a rocky substrate.
This is generally formed of calcrete (over
Pleistocene deposits or Cretaceous chalk), or
Tertiary Lamont Sandstone, or lithified Pleisto-
cene beach ridges. Where stromatolites have

15

Journal of the Royal Society of Western Australia, Vol. 62, Parts 1-4, 1979.

been localized by beach ridges, they occur in
curved bands which are often approximately
parallel to the present coastline.

Playford and Cockbain (1976) confirmed that
individual stromatolites are generally oriented
in the direction of wave movement, perpendi-
cular to the shore. However, they also demon-
strated that in some areas rows of stromatolite
ridges have grown parallel to the prevailing
wind and that the “leaning” intertidal stroma-
tolites lean towards the wind rather than in the
direction of wave movement. It appears that
the stromatolite ridges (here termed “seif
stromatolites”) have formed in response to
helical water vortices induced in shallow water
by strong prevailing southerly winds. These
ridges exhibit “tuning fork junctions” opening
upwind, and in this respect they resemble sub-
aerial seif dunes (which are thought to have
resulted from helical air vortices). However,
the mechanism whereby the prevailing wind
controls the inclination of “leaning” stromato-
lites is not fully understood.

Lithification of the Hamelin Pool stromatolites
is occurring in both intertidal and subtidal
environments, contrary to the suggestion of
Logan (1961) that exposure in the intertidal zone
is required for the induration of stromatolites.
The particulate sediment that has been trapped
and bound by algae is being cemented a few
millimetres or centimetres below the surface by
microcrystalline aragonite. Detailed studies have
not yet been carried out on the cementing
process to determine whether or not it is wholly
or partly biochemical, but it seems likely that
algal or bacterial action is at least partly
responsible for the aragonite precipitation.

Stromatolites at Hamelin Pool commonly form
linear reefs controlled by the existing shoreline
or by indurated submerged beach ridges. These
reefs are rigid and wave resistant but they lack
the skeletal framework required by many reef
definitions, having been built by non-skeletal
algae. The Hamelin Pool occurrences clearly
demonstrate the way in which stromatolites and
other cryptalgal bodies, when subject to early
cementation, can form rigid wave-resistant
bodies, and there is no good reason why these
should not be considered as representing a type
of organic reef. Many Proterozoic reefs and
some Palaeozoic reefs were apparently formed
in this way.

Growth rates

Growth-rate studies on intertidal and subtidal
stromatolites at Hamelin Pool show that the
living forms grow very slowly. The maximum
rate of growth observed is less than 1 mm per
year, and many forms appear to have virtually
reached a state of equilibrium, with growth
approximately balanced by erosion (Playford and
Cockbain 1976). It is likely that many of the
living stromatolites are many hundreds of years
old.

Tracks cut through the stromatolites by horse-
and camel-drawn wagons (when goods were
transported to and from stations around Hamelin
Pool by lighter) are still clearly defined at

several localities, little or no regrowth of stroma-
tolites having occurred in the tracks since they
were last used during the 1930s. This empha-
sizes the susceptibility of the Hamelin Pool
stromato'-ites to damage by human activities
(Department of Environmental Protection 1975,
Playford 1976a).

Classification

A geometric system of stromatolite classifica-
tion, based on the arrangement of crude
hemispheroids and spheroids, was proposed by
Logan et al. (1964), using the Hamelin Pool
stromatolites as a principal example. This
classification has been widely used, especially
among sedimentologists. However, it does not
take into account the internal fabric and micro-
structure of stromatolites, and is regarded as
inadequate for detailed studies by many workers
(Walter 1972).

Modern stromatolite specialists who have
adopted a biostratigraphic approach to stroma-
tolite research commonly use the Russian
binomial taxonomy of “group” and “form” in
classifying fossil stromatolites. As yet no one
has attempted to apply this classification to
the Hamelin Pool stromatolites. However, as
pointed out by Playford and Cockbain (1976),
they could form an excellent basis for testing
principles of stromatolite classification, as the
diverse living stromatolites have been built by
several mat types and extend through a range of
environments. If the taxonomic principles
applied to fossil stromatolites have any biological
validity they should also be applicable to the
modern Hamelin Pool stromatolites.

Hamelin Pool stromatolites : is the present the
key to the past?

The barred basin forming Hamelin Pool
represents an unusual environment in modern
seas, and the unique occurrence there of flourish-
ing stromatolites is linked to the hypersalinity
of this environment. Hypersalinity has caused
a major reduction in the number of metazoan
species in Hamelin Pool compared with adjoining
less-restricted areas of Shark Bay. A few
euryhaline species (such as the bivalve Fragum
erugatum and various fish) are abundant in
Hamelin Pool, but it is clear that algae grazers
(especially gastropods) are much reduced. This
allows stromatolites and associated algal mats to
flourish in Hamelin Pool, whereas elsewhere, in
waters of more normal oceanic salinity, they
tend to be “nipped in the bud” by metazoan
grazers.

The reduction in algae-consuming organisms
in Hamelin Pool resembles the situation that
prevailed in the world’s oceans during the
Proterozoic, when few metazoans had developed,
allowing stromatolites to develop widely.
Stromatolites subsequently declined progressively
during the Phanerozoic, as animal life diversified
and became more abundant, until today Hamelin
Pool is the principal remaining stronghold of
marine shallow- water stromatolites in the world.

16

Journal of the Royal Society of Western Australia, Vol. 62, Parts 1-4, 1979.

The history of stromatolite research at
Hamelin Pool illustrates the dangers inherent
in uncritical acceptance of modern environments
as definitive guides to the past, especially when
those environments are themselves imperfectly
known. Uniformitarian application of the
original Hamelin Pool stromatolite model led to
the widespread but mistaken belief that ancient
stromatolites are strictly intertidal phenomena
and their morphology is controlled solely by
environmental factors.

Apart from the fact that the original Hamelin
Pool observations were inaccurate, there can be
no justification for assuming that all ancient
stromatolites have formed under conditions
resembling those in Hamelin Pool today. The
factor of prime importance in development of
the Hamelin Pool stromatolites is hypersalinity,
and although some ancient stromatolites can be
shown to have formed in hypersaline environ-
ments, this was not the case for most ancient
stromatolites.

This is not to say that the Hamelin Pool
stromatolites are of no value in understanding
ancient stromatolites. Quite to the contrary,
they have much to teach us about stromatolite
growth mechanisms and biological and environ-
mental controls on stromatolite morphology.
This information is an important aid in inter-
preting ancient stromatolites, but what needs to
be emphasized is that evidence for the recon-
struction of ancient environments must be
sought primarily in the ancient rocks themselves.
Environments that favoured stromatolites in the
past do not favour them today; the present is
thus only partly a key to the past.

Devonian stromatolites in the Canning Basin

Introduction

Algal stromatolites are important constituents
of the Devonian reef complexes of the Canning
Basin. These complexes form a barrier-reef
belt which extends for about 300 km along the
northern margin of the basin, adjoining the
Precambrian Kimberley Block. They range
from Middle to Late Devonian in age.

Three basic facies are recognized in the com-
plexes: platform, marginal-slope, and basin

facies. There are also several named sub-facies,
the most important being the reef -margin, reef-
flat, back-reef, reefal-slope, and fore-reef sub-
facies (Playford 1976b).

The platforms were for the most part built
by stromatoporoids, corals, and algae in the
Givetian and Frasnian, and by algae in the
Famennian. The platform deposits accumulated
in near-horizontal beds, commonly with a
massive or crudely bedded reef margin. The
platforms stood some tens to hundreds of metres
above the surrounding inter-reef basins, and
were flanked by steeply dipping marginal-slope
deposits composed largely of platform-derived
debris, with contributions from indigenous
organisms and terrigenous sources. Depositional
dips in these deposits were commonly up to
35-40° in loose sediments, and up to vertical
where algal binding and precipitation occurred
together with early lithification. At the foot of

the slopes the marginal-slope deposits inter-
fingered with the flat-lying basin deposits, com-
posed largely of terrigenous material.

Stromatolites occur in the reef complexes in
the platform and marginal-slone facies (Playford
and Cockbain 1969; Playford 1973; Playford et ah
1976). In a few areas they extend from the
marginal-slope deposits into the adjoining part
of the basin facies. Those that occur in the
platform facies are believed to have grown in
shallow water (less than about 5 m in depth) ,
while the others have grown in relatively deep
water (to depths of 100m or more).

Shallow-water stromatolites

Shallow-water stromatolites and other crypt-
algal limestones are widespread in the platform
facies of the reef complexes, where they are
commonly the principal rock builders, especially
in the younger Frasnian and Famennian com-
plexes. Columnar forms occur most frequently
in the reef-margin sub-facies, and flat-bedded
algal-mat deposits and oncolites are common in
the back-reef sub-facies. Fenestral fabrics are
characteristic of the shallow-water stromatolites
and other cryptalgal deposits of the platform
facies, but are absent in the deep-water stroma-
tolites.

The columnar stromatolites of the platform
facies closely resemble the modern Hamelin Pool
stromatolites in both external morphology and
internal fabric. The close similarity between
the fenestral fabrics of these Devonian and
modern forms is very striking (Playford et al.
1976). It is believed likely that such fenestral
fabrics are characteristic of shallow-water
stromatolites, both modern and ancient.

Deep-water stromatolites

Stromatolites are conspicuous features of some
parts of the marginal-slope facies, extending
into adjoining parts of the basin facies, and
they also occur capping drowned reefs and
allochthonous reef blocks.

The marginal-slope stromatolites grew on
slopes with original inclinations ranging from a
few degrees up to near vertical. Stroma tolitic
algae are believed to have been responsible for
maintaining those depositional slopes that
exceeded the angle of rest (35-40°) for loose
debris. Both skeletal and non-skeletal algae
were involved in the construction of the stroma-
tolites, recognizable forms including Sphaero-
codium, Renalcis, Girvanella, and Frutexites
(although some authorities regard Renalcis as a
foraminifer rather than an alga).

The deep-water stromatolites are considerably
more diverse than the shallow-water forms in
the reef complexes. Playford et al. (1976) recog-
nized spaced columnar, contiguous columnar,
branching columnar, longitudinal, scallopeu,
reticulate, undulous, and domal deep-water
forms. In addition, large stromatolite bioherms
up to 1 km across are developed in some areas,
overlying drowned stromatoporoid-algal pinnacle
reefs. The most impressive of these occur near
Elimberrie Spring in the Oscar Range. Similar

82710 ( 2 )

17

Journal of the Royal Society of Western Australia, Vol. 62, Parts 1-4, 1979.

stromatolite cappings, but on a much smaller
scale, occur on top of allochthonous reef blocks
(derived from platform margins) in the margin-
al-slope facies.

Depth relationships of the deep-water stroma-
tolites have been deduced from palaeobathy-
metric reconstructions. The depositional and
post-depositional components of an observed dip
in marginal-slope deposits can be determined
using geopetal fabrics (Playford et al. 1976),
and on this basis palaeobathymetric measure-
ments can be made. Thus, where an observed
dip in marginal-slope deposits is solely deposi-
tional the difference in water depth between any
two points on a bedding plane is the same as
the present elevation difference between them.
Appropriate corrections are made where post-
depositional tilting is involved. This method
has been applied at several localities where
stromatolites occur in situ in marginal-slope
deposits, and reconstructions indicate that some
of the deep-water stromatolites must have grown
in water as deep as 100 m or more.

Well-developed deep-water stromatolites are
associated with strongly condensed sequences
(i.e. very slow deposition). They grew on those
areas of the marginal slopes that were receiving
very little sediment and on bare elevated features
such as drowned pinnacle reefs and alloch-
thonous reef blocks. They are best developed
in the Frasnian-Famennian Virgin Hills and
Napier Formations, especially in the early
Famennian parts. Evidence for the condensed
nature of the stromatolite-bearing horizons is
provided by conodont zones, which are very much
thinner than in equivalent sections lacking
stromatolites. It is deduced that the stromato-
lites grew very slowly; the average annual
growth rate may commonly have been as low
as 2 fim.

Many of the deep-water stromatolites in the
Canning Basin reef complexes are coloured
bright red, due to the presence in them of finely
divided iron oxide (mainly hematite). In some
stromatolites there is a pronounced concentration
of iron oxide in and around algal filaments,
especially those of the genus Frutexites, in which
the iron content may be as high as 30%. Other

algae which commonly show iron concentration
are Girvanella and (to a lesser extent) Sphaero-
codium. Microprobe analysis indicates that low
concentrations of manganese also occur with
the iron-rich filaments. It is believed that
bacteria living in association with the algae are
most likely to have been responsible for this iron
and manganese deposition.

The columnar deep-water stromatolites com-
monly grew approximately vertically, either
towards the light (phototropic) or under the
influence of gravity (geotropic). It is not known
whether all the stromatolitic organisms were
photosynthetic; some stromatolites could have
been formed by heterotrophic algae or bacteria.

The deep-water stromatolites grew as hard
and rigid bodies; many are encrusted by crinoid
and coral holdfasts, and some show evidence of
early fracturing, the fractures being filled with
early submarine cement or sediment, forming
neptunian dykes. Some of the stromatolites
have formed primarily through the trapping
and binding of clastic particles (including terri-
genous detritus) by non-skeletal algae (and/or
bacteria) associated with biochemical or
inorganic precipitation of cement. Others con-
tain little or no clastic material and such
stromatolites were formed primarily or wholly
by algal precipitation of carbonate.

Apart from the sessile crinoids and corals
mentioned above, the fauna of the condensed
sequences associated with the stromatolites is
characterized by open-marine pelagic organisms:
conodonts, ammonoids, and nautiloids. These
fossils are often very abundant, especially in the
interareas between stromatolites.

Characteristics of shallow-water and
deep-water stromatolites

The characteristics of deep-water stromatolites
in the Canning Basin reef complexes are
distinctly different from those of the shallow-
water forms. These differences are summarized
in Table 1. It is suggested that many of these
features may be applicable in other parts of
the stratigraphic record for distinguishing
between shallow- and deep-water stromatolites.

Table 1

Characteristics of deep-water and shallow -water stromatolites in the Canning Basin

Deep-water stromatolites

Shallow-water stromatolites

Fenestral fabrics absent
Usually finely laminated
Diverse assemblage of forms

In condensed sequences

Grew on depositional slopes, drowned reefs, and
allochthonous blocks

Pelagic faunas common

Some forms encrusted with crinoids and corals

Iron and some manganese precipitation important
in certain forms

Commonly red or reddish brown

Fenestral fabrics common

Usually weakly laminated or unlaminated

Columnar forms only, associated with oncolitic and
fenestral limestones

Not in condensed sequences

Grew on near-horizontal limestone platforms

Associated with reefal and biostromal organisms,
especially stromatoporoids and Renalcis

Not encrusted by corals or crinoids
No significant iron or manganese precipitation

Commonly white or pale yellow

18

Journal of the Royal Society of Western Australia, Vol. 62, Parts 1-4, 1979.

The deep-water stromatolites show some
resemblance to ferromanganese deep-water bac-
terial stromatolites known from modern seas
(Monty 1973). Both occur in condensed
sequences, are generally finely laminated, lack
fenestral fabrics, and are associated with pelagic
faunas. Iron and manganese concentrations
also occur in both, but these concentrations are
much higher in the modern ferromanganese
stromatolites.

Proterozoic stromatolites

During the late 1960s the University of
Adelaide became the main centre of Precambrian
stromatolite research in Australia, under the
direction of M. F. Glaessner. He introduced
principles of stromatolite classification and
methods of stromatolite reconstruction based on
serial sectioning that had been applied for some
time in the Soviet Union, where stromatolites
had been used extensively for Precambrian
correlations. Two of his students, M. R. Walter
and W. V. Preiss, began a wide-ranging study
of Australian Precambrian stromatolites, which
is still continuing. Their two principal publica-
tions which deal with Western Australian forms
are by Walter (1972) and Preiss (1976). K.
Grey of the Geological Survey of Western
Australia is also conducting research on Pre-
cambrian stromatolites from this State.

Walter (1972) described stromatolites from the
Early Proterozoic Fortescue Group (ca. 2 400
m.y.) of the Hamersley Basin, the Early
Proterozoic Wyloo Group (ca. 1 900 m.y.) of the
Ashburton Trough, the Middle Proterozoic
Bangemall Group (ca. 1 000 m.y.) of the Bange-
mall Basin, and the latest Proterozoic or earliest
Cambrian Antrim Plateau Volcanics of the Ord
Basin.

In addition to these Western Australian
occurrences Walter also described Proterozoic
stromatolites from the Amadeus, McArthur and
Georgina Basins of the Northern Territory and
Queensland. His results suggested that stroma-
tolites could be used for inter-regional correla-
tion within Australia, but he noted some
anomalies in correlations with the Russian
sequence. One of the stromatolite groups
(Patomia) he recorded in the Early Proterozoic
Wyloo Group is indistinguishable from a Late
Proterozoic (Vendian) form in the Soviet Union.

Preiss (1976) described an assemblage of
Proterozoic stromatolites from the Nabberu and
Officer Basins in Western Australia. The
Nabberu Basin occurrences are in the Early
Proterozoic Earaheedy Group (at least 1 700
m.y.), while those from the Officer Basin are
from Woolnough Hills and “Central Neale” (near
Yeo Lake) in rocks which may correlate with
the lima Beds (probable Late Proterozoic).

The stromatolite groups recorded from the
Officer Basin are consistent with the expected
Late Proterozoic age, based on correlation with
other stromatolite occurrences of this age in
South Australia. However, the Early Proterozoic
Nabberu Basin stromatolites are anomalous, as
the stromatolite groups present have only previ-
ously been recorded elsewhere from Late Protero-

zoic rocks — Minjaria from the Late Riphean,
and Tungussia from the Middle Riphean to
Vendian of the Soviet Union. Kulparia, another
form which may be present (recorded as ?cf.
Kulparia ), is known elsewhere in Australia from
Late Riphean or Vendian equivalents. It there-
fore seems that the time range of these stroma-
tolite groups is considerably longer than had
previously been believed, assuming that the 1 700
m.y. dating (K/Ar based on glauconite) is
correct.

Consequently, Preiss (1976, 1977) advocates
caution in dating Precambrian sediments on the
basis of stromatolites alone, especially where
correlations are made using stromatolite groups
rather than forms. Although the Russian bio-
stratigraphic scheme based on stromatolites
appears to have validity throughout the Soviet
Union, detailed stromatolite studies are still in
their infancy in most other countries, and it
remains to be seen how much of the Russian
biostratigraphy can be applied on an inter-
continental scale.

Acknowledgements . — This paper is published with the
permission of the Director, Geological Survey of Western
Australia. A. E. Ccckbain and M. R. Walter are thanked
for their comments on the draft manuscript.

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20

Journal of the Royal Society of Western Australia, Vol. 62, Parts 1-4, 1979, p. 21-31.

Aspects of Science in Western Australia 1829-1979.

A volume to commemorate the 150th Anniversary of the founding of Western Australia.

The contribution of geochronology to Precambrian studies in

Western Australia

by J. R. de Laeter and A. F. Trendall

Physics Department, Western Australian Institute of Technology, South Bentley, W.A. 6102
Geological Survey of Western Australia, 66 Adelaide Terrace, Perth, W.A. 6000

Abstract

In 1906, one decade after the discovery of radioactivity, Rutherford demon-
strated that the accumulation of radiogenic daughter products in rocks and minerals
containing radioactive elements could be used to calculate their ages. This event
initiated the discipline of physical geochronology, whose development and application,
outlined in this paper, gradually enabled a firm time scale to be established
for the study of earth history. Such a scale had a particularly powerful effect on
the study of Precambrian rocks, for which no other effective internal age criteria
had been available. The U-Pb, Th-Pb, K-Ar, and Rb-Sr methods were successively
developed and refined to produce compatible results of great precision; the mass
spectrometer played a key part in this process. In Western Australia, an attempt
was made to obtain a mineral U-He age as early as 1910, and data obtained at
that time were used in 1926 to calculate U-Pb and Th-Pb ages. But the work was
not pursued, and geologists were slow to realise the potential of the new discipline;
it was not until the early 1960s, some 10 years after the establishment of the first
geochronological laboratory in the State, in the Physics Department of the University
of Western Australia, that geologists working on the Precambrian came to appreciate
the full value of physical age determinations for their work. Since that time,
continuing work at the Australian National University in Canberra, and at the
Western Australian Institute of Technology, has combined with the more recent work
of other groups to produce a good understanding of the broad geochronological
framework of the State. Nevertheless, many problems still remain, but newly
developing methods show promise for their resolution.

Introduction

With the benefit of hindsight it is clear that
the introduction, during the first decade of this
century, of rock and mineral age determination
by methods based on radioactive decay was an
event of crucial significance for geology, in that
it initiated the systematic constraint of his-
torical geology within a reliable time scale.
However, to contemporary geologists the impor-
tance of this event was by no means self-evident,
and the development of physical geochronology
in relation to ‘classical’ geology deserves wide
attention as an example of the complex processes
at work in the progress of scientific ideas.

For Western Australia the subject has special
interest, both through the importance of geo-
chronology in a State with such a vast area of
exposed Precambrian rocks of different ages, and
also because certain important conceptual
advances in geochronology were made in Perth.
In outlining, in a paper of this length, the
progress of geochronology in Western Australia
and its impact on Precambrian studies, we have
had to assume that the reader has an acquaint-
ance with both the geology and geography of
the State. Readers lacking this, but neverthe-

less interested in detailed studies, will find
adequate information in both areas in Memoir
No. 2 of the Geological Survey (G.S.W.A. 1975);
readers of this paper who wish to extend their
knowledge of the development of geochronology
will find a book compiled by C. T. Harper (1973)
of great interest.

Critical events in geochronology

Discovery of radioactivity

Early attempts to measure geological time were
based on such time-dependent phenomena as
rates of sedimentation, the rate of cooling of
the earth, and the accumulation of sodium in
sea water. All these techniques were quantita-
tively crude, and involved sweeping assumptions,
and it was apparent that a new approach to
the problem was required. This came about in
a curious fashion. In 1896 Henri Becquerel, a
French physicist, found that uranium salts
placed near a photographic plate in a drawer
in his work table had produced an image of
a key which was accidently lying on the plate.
He realised that the uranium was emitting
invisible but penetrating rays — the element was
in fact “radioactive”.

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Journal of the Royal Society of Western Australia, Vol. 62, Parts 1-4, 1979.

Rutherford and Soddy, investigating the
phenomenon discovered by Becquerel, found that
the empirical facts of radioactivity could be
explained by assuming that radioactive atoms
disintegrated at characteristic rates to form new
atoms of other elements. The rate of decay of
a radioactive parent to a stable daughter is
proportional to the number of parent atoms, N,
present. This law of radioactive decay can be
expressed as

-Xt

N = N 0 e

where N 0 is the number of radioactive parent
atoms present at the time of formation of a
mineral or rock, t measures the time elapsed
since their formation and X is the decay con-
stant of the radioactive nuclide. It was shown
that the decay constant did not vary over
extreme variations in the physical environment
and has remained unchanged at least over
geological time periods.

Chemical methods in geochronology

Lord Rutherford quickly realised that the
accumulation of stable radiogenic daughter
products, such as helium and lead in uranium
bearing minerals, could be used to measure
geological time (Rutherford 1906). Unfortun-
ately the only means available at that time to
measure helium and lead were chemical tech-
niques, and both these methods had serious
shortcomings.

Strutt (1910) showed that helium leaked from
uranium-rich ores, even under laboratory con-
ditions, and thus the measured helium ages could
only be regarded as minimum estimates. As
a result the helium method fell into disuse for
nearly 20 years, until Dubey and Holmes (1929)
showed that the method could be used to
measure the age of common rocks.

After some promising early work on the
uranium-lead method by Boltwood (1907) and
Holmes (1911), it was realised that radiogenic
lead was produced by both uranium and thorium,
and since these two elements often occurred
together in minerals, it was impossible to disen-
tangle the two decay chains by purely chemical
methods. Another problem was the presence
of common lead, included in minerals at the
time of their formation, which could not be
distinguished from the radiogenic lead except
by using corrections based on atomic weight
determinations. Thus the chemical lead method
could only be used for rich uranium-bearing
ores and was therefore not applicable to common
rocks where the amount of uranium was exceed-
ingly small.

One of the first earth scientists to appreciate
the importance of radiometric-based geochron-
ology was Arthur Holmes. He was one of the
early proponents of the uranium-lead method,
and urged the establishment of a radiometric
time scale, and the application of radiometric
dating to the crystalline basement rocks of the
Precambrian. His book entitled “The Age of
the Earth” (Holmes 1927), did much to publicise
the emerging field of geochronology, particularly
among geologists.

Physical methods in geochronology

A major breakthrough in the extension of
geochronology came from a most unexpected
direction. F. W. Aston, working almost in isola-
tion at the Cavendish Laboratory, developed a
mass spectrograph employing both electric and
magnetic fields, which was far more sensitive
than earlier versions. By studying the isotopic
composition of lead he observed isotopes at
masses 206, 207 and 208 (Aston 1927). Aston
then analysed a uranium-rich sample of brog-
gerite and showed that it was highly enriched
in lead 206 (Aston 1929). The age reported for
this mineral heralded a new era in geochron-
ology, in that this age was based on an isotopic
rather than a chemical analysis.

Since that time the mass spectrometer has
become the tool of every practising geochron-
ologist — a veritable time machine for the
exploration of the past (Harper 1973). Between
1929 and 1941 mass spectrometric techniques
developed rapidly, mainly as a result of the work
of Alfred Nier at the University of Minnesota.
Nier et al. (1941) reported the results of isotopic
analyses of lead separated from both radioactive
and lead ore minerals. The age of one monazite
sample gave a 207 Pb/ 206 Pb age of 2 570 ± 70 m.y.,
which was the first substantial isotopic evidence
of the immense age of the Precambrian. Nier
was also instrumental in determining the isotopic
composition of uranium (which was a vital
factor in atomic energy research), in discovering
the geochronologically important isotope 40 K, and
showing that radiogenic 40 Ar could accumulate
in potassium-rich minerals (Aldrich and Nier
1948). Nier has rightly been called the “father
of modern mass spectrometry”.

The emergence of new techniques in
geochronology

After a bright beginning early in the present
century, geochronology developed slowly. The
late 1940s and 1950s however, saw an enormous
burst of activity which established the feasibility
of the potassium-argon and rubidium-strontium
techniques, and extended the traditional U-Pb,
Th-Pb method. After World War II isotopically
enriched tracers became available, and coupled
with further advances in mass spectrometry,
enabled very low concentrations of uranium,
thorium and lead to be accurately measured.
This method of analysis, known as stable isotope
dilution, has recently been described by de Laeter
and Rosman (1977).

It was now possible to obtain three indepen-
dent ages from the U, Th-Pb system: 206 Pb/ 238 U,

207p b/ 23 5U 0r 2<>7p b/ 20 6pb and 208p b/ 232 Th Wethei'ill

(1956) used a ratio plot of 206 pb/ 238 U against
207 Pb/ 235 U to show that a phase which has
remained a closed system to uranium and lead
will have 20(i pb/ 238 u ages equal to 207 pb/ 235 U ages,
and that the data for systems of different age
will lie along a curved line called “concordia”.
Phases having the same age, and subject to lead
loss or uranium gain will, under prescribed
conditions, have data that lie along a straight
line which intersects concordia. The lower
intersection may represent the time of the

22

Journal of the Royal Society of Western Australia, Vol. 62, Parts 1-4, 1979.

episodic event and the upper intersection the
age of the phase itself. The work of Silver and
Deutsch (1963) showed that U-Pb zircon analyses
could be used to give the time of original crystal-
lization of metamorphosed Precambrian cratonic
areas.

The decay of the radioisotope 40 K to 40 Ar forms
the basis of a dating technique which can be
applied to rocks and minerals between a few
tens of thousand and several billion years old —
a broader range than can be determined with
other radioactive methods. Because potassium
is a common rock-forming mineral and argon is
retained in many minerals over geologically
long periods of time, most rock types can be
successfully analysed by this technique. After
the early work by Aldrich and Nier (1948),
Reynolds (1956) developed a high-sensitivity
mass spectrometer for noble-gas analysis, and a
number of studies determined the argon retenti-
vities of a wide variety of geological materials.
Application of the K-Ar method to young vol-
canic rocks has been described by McDougall
(1966), whilst the technique has also been used
successfully to date the times of reversals of
the earth’s magnetic field (see for example
Cox et al. 1968).

However, the K-Ar method had serious short-
comings, most of which were related to the
migration of argon in mineral assemblies.
Fortunately, a variation of the K-Ar method
described by Merrihue and Turner (1966), has
enabled the imperfect retention of radiogenic
argon, or the presence of excess 40 Ar to be
measured. This technique is known as the “ 39 Ar
method”, as it involves the conversion, by irradi-
ation with fast neutrons, of a proportion of the
39 K nuclei to 39 Ar.

Another new technique that emerged at about
the same time was based on the decay of 87 Rb
to 87 Sr (Hahn and Walling 1938). Eleven years
were to elapse before Ahrens (1949) published
the results of an extensive investigation into
Rb-Sr dating based on a spectrochemical rather
than a mass spectrometric technique, and showed
that it was possible to obtain ages by the Rb-Sr
method.

The problem of separating the very small
amount of rubidium and strontium encountered
in common rock forming minerals in a form
pure enough to analyse in a mass spectrometer
was overcome by the use of ion-exchange
chemistry (Aldrich et al. 1953), enabling common
strontium to be distinguished from radiogenic
87 Sr, and accurate mass spectrometric results to
be obtained. Throughout the 1950s Rb-Sr dating
was applied to micas and feldspars from a
number of rocks, and the method gained
increasing recognition. Schreiner (1958) then
showed that whole rock samples could be used
for Rb-Sr dating in addition to mineral
separates.

This observation led to a major advance in
Rb-Sr geochronology, in that Compston and
Jeffery (1959) were able to show that minerals
could lose radiogenic 87 Sr some time after crystal-
lisation, but that the radiogenic 87 Sr was still
retained by the whole rock itself. The Compston-

Jeffery model provided an explanation for
discordant Rb-Sr ages and showed that primary
crystallisation ages could be determined in
addition to metamorphic events in isotopically
disturbed systems. Further details of this work
are given later in this paper. Nicolaysen (1961)
used a simplified graphical method by plotting
87 Sr/ 86 Sr against 87 Rb/ 86 Sr in which members of
a comagmatic suite of igneous rocks which have
remained chemically closed to rubidium and
strontium form a straight line called an “iso-
chron”, the slope of which is related directly to
the age of the rocks, while the intercept on the
87 Sr/ 86 Sr axis defines the value of this ratio in
the magma source region at the time of differen-
tiation. The Rb-Sr method has been used more
extensively in Western Australia than any other
technique.

The impact of modern technology

In the early 1960s geochronology was hampered
by poor analytical precision, and the resulting
interlaboratory discrepancies made it difficult
to compare results on an international basis.
As a result of these problems a number of inter-
national standards were produced to assist
laboratories in producing data of high quality.
For example, the Eimer and Amend standard
was distributed widely to laboratories involved
with Rb-Sr analyses, and more recently the
National Bureau of Standards have produced
stoichiometric salts of rubidium and strontium
of known isotopic composition. Standard silicate
rocks have also been used by many laboratories
to check their elemental abundance measure-
ments, so that geochronological data can now be
compared with greater reliability than in the
past.

Commercial mass spectrometers have also
improved considerably over the past decade.
Technological advances including solid-state
electronics, peak switching, on-line data reduc-
tion by minicomputers and high- vacuum tech-
nology, have greatly improved the precision and
speed at which isotopic ratios can be measured,
and expanded the range of geological samples
that can be used. Improvements in ion-
exchange chemistry and the use of low contam-
ination “clean rooms” have also played a major
role in improving the quality of the data, and
have allowed problems to be successfully tackled
which would have been impossible even ten
years ago. One of the factors in this techno-
logical revolution has been the Apollo pro-
gramme. Considerable information concerning
lunar chronology has been accumulated by a
variety of geochronological techniques, in which
technical problems of considerable magnitude
have had to be overcome. The experience so
gained has been of inestimable value to terres-
trial geochronology. A further improvement in
the comparison and assessment of geochrono-
logical data has resulted from new measurements
of the decay constants of 235 U and 238 U ( Jaffey
et al. 1971) and the adoption of a new value of
1.42 x HT 11 yr’ 1 for 87 Rb (Steiger and Jager
1977).

23

Journal of the Royal Society of Western Australia, Vol. 62, Parts 1-4, 1979.

Western Australian geochronology

Early ages derived from Simpson’s work

The earliest attempt to determine the age of
a Western Australian mineral by a method based
on radioactive decay was made by E. S. Simpson,
then Chemist and Assayer of the Geological
Survey of Western Australia, in 1910 (Simpson
1910, 1911, 1912). Simpson had received, in that
year, samples of a bright-yellow ochreous
uranium mineral, which he named pilbarite,
from a pegmatite at Wodgina, in the Pilbara
Block. As he was acquainted with the work of
Rutherford and Strutt on the use of helium in
age determination, he carefully analysed the
material for helium, and calculated an age of
13m.y.; but he pointed out that as it was a
secondary weathering product, and in any case
may also have lost radiogenic helium, this age
probably had no significance.

At about the same time Simpson also fully
analysed two associated uranium minerals from
the same pegmatite, which he identified as
mackintoshite and thorogummite, parental to
the pilbarite. An intriguing question then
presents itself as to why Simpson failed to
calculate the ages of all three minerals from
the amount of lead present. He had determined
this, and had specifically commented on its
derivation by radioactive decay; he was also
generally familiar with the work of Boltwood,
for he mentions him by name (Simpson 1912),
though he does not refer to Boltwood’s 1907
paper.

Whatever the reason for this puzzling omission,
fifteen years went by before L. A. Cotton,
Professor of Geology at the University of Sydney,
used Simpson’s published analytical results to
calculate chemical lead ages for these minerals,
and also for fergusonite from another Pilbara
pegmatite at Cooglegong, an analysis of which
had by then been published by Simpson (1919).
Cotton’s (1926) calculated ages of 620, 1 475,
1 460 and 3 840 m.y. respectively for fergusonite,
mackintoshite, thorogummite, and pilbarite took
account of the lead contribution by thorium.
Holmes and Lawson (1927) independently
revised Cotton’s calculations, and the inclusion
of the age of “mackintoshite, etc” as 1 260 m.y.
in Holmes’ (1927) popular book “The age of the
Earth”, in which it was the oldest age recorded,
placed the significance of Simpson’s 1910 ana-
lytical work before a wide audience.

It now appears that this age is too low by a
factor of at least two, and this makes it even
more ironic that it was probably largely due to
Holmes’ book that a widespread belief arose that
the Precambrian of Western Australia contained
the most ancient rocks of the Earth’s crust.

Miscellaneous later results

Simpson did not maintain his interest in
mineral age determination, and in a letter of
1927 on Government Chemical Laboratories files
he politely rejected a suggestion from Sir
Douglas Mawson that he should recommence
such work. Chemical lead age determinations
were later carried out, in 1953, by the Chemical

Laboratories on two samples of allanite, collected
by A. F. Wilson from pegmatites in the Fraser
Range and at Doubtful Island Bay (Prider 1955).
The ages, of 1210 ± 50 and 1 390 ± 50 m.y.,
are in good accord with subsequent K-Ar and
Rb-Sr work on biotite (Aldrich et al. 1959), as
well as with more recent Rb-Sr results (Bunting
et al. 1976).

These isolated results were not part of a
continuing programme, and the same comment
applies to a group of ages estimated from the
isotopic compositions of galena samples supplied
by J. D. Campbell and R. T. Prider to J. Tuzo
Wilson, of the University of Toronto, in 1953
(Prider 1955). These were the first results for
any Western Australian minerals by a method
employing mass spectrometry.

Establishment of systematic work in the 1950s

In 1950, on his way to take up an appointment
at the Australian National University, Professor
(later Sir) Marcus Oliphant visited the Physics
Department of the University of Western
Australia, and spoke of the need for an Austra-
lian research programme in geochronology,
particularly in view of the widespread belief
that the continent contained the Earth’s oldest
rocks (Jeffery 1976). Oliphant had worked with
Lord Rutherford in Cambridge, and was familiar
with the physical techniques used in geochrono-
logy.

As a result, two younger staff members, P. M.
Jeffery and A. H. Morton, terminated the con-
struction of a small accelerator and began
instead to build a mass spectrometer, intended
ultimately for a continuing geochronological
programme. Jeffery (1976) has given a graphic
account of the difficulties involved. The instru-
ment first operated in late 1952, but its perform-
ance was not satisfactory, and during 1953 it was
decided to abandon the project unless more
financial support could be found. Fortunately,
a grant became available from the Carnegie
Institute of Washington, sufficient not only to
bring the home-made instrument into effective
operation but also to provide a second, new,
Nier-type mass spectrometer. It was on this
instrument, commissioned by D. Greenhalgh,
that the first isotopic U-Pb age was obtained
from a Western Australian mineral. The tanteu-
xenite used came from a pegmatite at Wood-
stock, again in the Pilbara Block, and yielded a
maximum age of 2 790 ± 25 m.y. (Greenhalgh
and Jeffery 1959).

It was only after Jeffery had taken up a
Carnegie Fellowship at the Department of Ter-
restrial Magnetism in Washington in 1955 that
the potential of the Rb-Sr and K-Ar methods
was fully appreciated, and on his return to Perth
programmes in both these methods were begun.
Although a number of K-Ar determinations were
carried out on micas and feldspars from pegma-
tites and granites, the main achievements of
the group thus established lay in the Rb-Sr
method, in which work Jeffery was joined by
W. Compston and G. H. Riley.

Towards the end of the 1950s the validity of
the Rb-Sr method was being questioned, since

24

Journal of the Royal Society of Western Australia, Vol. 62, Parts 1-4, 1979.

mineral separates from whole rock samples were
giving discordant ages. A granite from the
Boy a quarry near Perth was analysed by Rb-Sr
geochronology, together with the biotite and
microcline separates. The resultant ages were
650 m.y. for the biotite, 1 290 m.y. for the micro-
cline and 2 430 m.y. for the whole rock
(Compston and Jeffery 1959). The authors
suggested that the mineral separates had lost a
significant proportion of their radiogenic Sr
some time after crystallisation, after which
metamorphic events caused the Rb-rich minerals
to lose radiogenic 87 Sr which was simply redis-
tributed within the whole rock system.

The model proposed by Compston and Jeffery
represented a most important advance in the
interpretation of discordant Rb-Sr ages and
established the Rb-Sr technique as a powerful
tool in geochronological studies of isotopically
disturbed systems (Harper 1973). Fairbairn et
al. (1961) applied the Compston- Jeffery model
to a number of discordant ages from Sudbury
in Canada, and concluded that “we therefore
regard the whole-rock approach as a milestone
in interpretation of discordant Rb-Sr data”.
The Compston-Jeffery model was elaborated in
a number of important publications (Compston
et al. 1960, Riley and Compston 1962, Riley 1961)
which set the seal on this contribution by the
laboratory.

An important geological contribution to the
programme was made by A. F. Wilson, of the
Geology Department of the University of
Western Australia. His energetic research
interests in the southern Yilgarn Block, and in
the Albany-Fraser Province, were a significant
control over the material selected for analysis.
The seminal 1960 paper by Wilson, Compston,
Jeffery and Riley (Wilson et al. 1960), in which
36 Rb-Sr ages (19 of them new), 16 K-Ar ages
(4 new), and 21 U-Pb ages were presented and
discussed together was the first systematic
attempt to relate these results to the tectonic
evolution of the Western Australian, and in this
case also the Australian, Precambrian. Even so,
their potential in this respect was restricted,
since apart from mineral ages from isolated
pegmatites of uncertain geological relationships
in the Halls Creek Province, the Pilbara Block,
the Northampton Block, and the Albany-Fraser
Province, the reported ages were confined to the
southern parts of the Yilgarn Block; the main
factor in the choice of the pegmatite minerals
was, necessarily at that stage, their chemical
suitability.

The changed situation of the 1960s

In 1959 it might have been supposed that all
was set for a major expansion of geochrono-
logical work at the University of Western Aus-
tralia, but in fact by 1961 such work had ceased
(Jeffery 1976). The prime reason for this was
probably a growing appreciation, from the
practical experience gained, that the establish-
ment and maintenance of a geochronological
laboratory of the size and scope that was needed
in Australia was, quite simply, an inappropriate
task for a comparatively small Physics

Department which wished also to retain a range
of representative research interests. The decision
to terminate such a successful research pro-
gramme was made easier by a concurrent move
by the Australian National University to estab-
lish a laboratory better equipped to supply
national needs in Canberra, in association with
the Commonwealth Bureau of Mineral Resources.
Compston moved to that laboratory in 1961,
while Riley took up a research fellowship at the
Australian Institute of Nuclear Science and
Engineering in New South Wales. Coinciden-
tally, Wilson, the major geological contributor
in the group, simultaneously moved to Queens-
land, and “a comprehensive geologic age
determination programme which is being
developed to outline the geochronology of the
Australian continent” (Greenhalgh and Jeffery
1959) was abruptly discontinued.

That Precambrian geochronological studies in
Western Australia were not disastrously set back
by this event is wholly due to the fact that,
from Canberra, Compston retained a strong
interest in Western Australian work. This was
effectively achieved by initiation of Ph.D. studies
in restricted areas, and theses which were com-
pleted or initiated in this way during the 1960s
include those of Turek (1966), Bofinger (1967),
Worden (1970), and Gray (1971). During this
period Compston maintained a good liaison with
the Geological Survey of Western Australia, and
some results were published in association with
it (Leggo et al. 1965). In respect to Western
Australian work, he also joined in geological
liaison with other university departments, and
encouraged such liaison among his students and
colleagues, so that consequent publications result
from a wide range of collaborative work.

Two characteristics differentiate the Western
Australian work carried out from Canberra in
the 1960s from the earlier Perth work. Firstly,
the necessity for close integration of field
geology and geochronological studies was
recognised. The work of Bofinger (1967) on the
eastern arm of the Halls Creek Province provides
a typical example of this: as a member of the
Bureau of Mineral Resources, Bofinger took part
in the joint field mapping with the Geological
Survey of Western Australia, and collected and
observed the geological relations of almost all of
the samples he analysed. Secondly, and as a
consequence of this recognition, an attempt was
made to extend geochronological work more
widely over the Western Australian Precambrian.
Thus from work during this period results were
obtained from the Kimberley region (Bofinger
1967, Bennett and Gellatly 1970), from the
Pilbara Block (Arriens 1967, Compston and
Arriens 1968), from the Hamersley Basin (Leggo
et al. 1965), from the Gascoyne Province
(Compston and Arriens 1968), from the Bange-
mall Basin (Compston and Arriens 1968), from
the Musgrave Block (Compston and Nesbitt
1967, Gray 1971), from the Albany-Fraser
Province (Arriens and Lambert 1969, Turek and
Stephenson 1966), from the Yilgarn Block
(Turek 1966, Worden 1970) and from ‘basement’
rocks encountered in petroleum exploration
drilling in the Officer Basin (Bofinger 1966), and

25

Journal of the Royal Society of Western Australia, Vol. 62, Parts 1-4, 1979.

the Canning Basin (Johnstone 1961). A note-
worthy early publication of this period recorded
a pioneer attempt to establish the depositional
age of a shale by the Rb-Sr method (Compston
and Pidgeon 1962).

Consolidation in the 1970s

It was this emphasis on the integration of
field geology and geochronology that brought
home to the bulk of geologists concerned with
the Western Australian Precambrian in the
1960s that isotope geochronology had become
an indispensable tool for the solution of many
problems, and a positive demand for results
began to grow. Two main factors made it seem
unlikely that Compston’s laboratory alone could
satisfy this. These were, firstly, the constraining
effects of the need to define most work in terms
of independent higher degree parcels, and an
increasing commitment of the laboratory to
prepare for and carry out work on returned
lunar samples, first received after the USA
Apollo 11 mission in 1969.

In response to this situation, we discussed,
early in 1968, as representatives of the Physics
Department of the Western Australian Institute
of Technology (JRdeL) and of the Geological
Survey of Western Australia (AFT), the possi-
bility of establishing a co-operative geochrono-
logy programme based on mass spectrometric
facilities already in operation at the Institute
of Technology. The ranges of geological and
physical expertise within the respective groups
appeared to be potentially complementary, and
it was agreed to proceed on a trial basis, with a
number of restricted projects.

For these initial projects rock bodies were
chosen for which clear-cut geological relation-
ships indicated the likelihood that they would
each yield a single age of known significance;
and a further restriction was applied that each
should be chemically amenable to a whole-rock
Rb-Sr isochron approach, using for each sample
a combination of X-ray fluorescence analysis of
the Rb/Sr ratio and a single unspiked mass
spectrometer run for the 87 Sr/ 86 Sr ratio. One
final constraint on project choice should not be
overlooked: close liaison was maintained with
the Australian National University geochrono-
logical laboratory to avoid duplication of work,
and this liaison was also crucial in the establish-
ment of experimental procedures.

The anticipated mutual advantages in
co-operation between the Geological Survey and
the Institute of Technology were amply con-
firmed in practice. The first results were pub-
lished in 1970 (de Laeter and Trendall 1970),
and a total of 19 papers have appeared from a
currently continuing programme. The basic
restriction to the Rb-Sr method has been main-
tained throughout, but no problem which could
be theoretically solved by the method is now
excluded by analytical constraints, although
regard in planning is paid to the desirability of
maintaining an acceptable ratio of geologically
meaningful results to invested analytical time.

Two important controls over the choice of
problem to attack have been the progress of the
Geological Survey regional mapping programme,
and the conscious attempt to identify priority
areas, in which work seems likely to yield the
most productive results. Work published so far
has been widely distributed over the State, and
includes results from the Pilbara Block (de
Laeter and Trendall 1970, de Laeter and Blockley
1972, de Laeter et al. 1975, Lewis et al. 1975, de
Laeter et al. 1977), from the Hamersley Basin
(de Laeter and Trendall 1971, Trendall and de
Laeter 1972, de Laeter et al. 1975. Hickman and
de Laeter 1977), from the Yilgarn Block
(Muhling and de Laeter 1971, Bunting et al.
1976), from the Paterson Province (Trendall
1974), from the Gascoyne Province (de Laeter
1976), from the Bangemall Basin (Gee et al.
1976) and from the Albany-Fraser Province
(Bunting et al. 1976).

Although we have chosen to regard the
re-establishment of geochronological work in
Perth as the distinguishing character of the
1970s we emphasize that there was no termina-
tion of Canberra work. This continued through
the 1970s with the same general policy and
objectives that had applied before. However,
a significant innovation during this period was
the initiation of Pb-Pb work on trace lead in
minerals (Oversby 1976), and subsequently of
the U-Pb method on extracted zircons. It is
a special objective of this work to identify the
oldest rocks of the Australian continent. The
3 450 m.y. age for dacite of the Duffer Forma-
tion, in the Pilbara Block, recently published
by Pidgeon (1978), is the oldest age so far
determined from the Archaean of Australia;
Pidgeon’s continuing work on zircons will be an
important addition to Western Australian geo-
chronology. Some work on Western Australian
material was initiated and carried out by the
Bureau of Mineral Resources in Canberra during
the 1970s (Page et al. 1976).

In addition to the continuing geochronological
programmes in Perth and Canberra, which we
have discussed in some detail, the 1970s saw the
initiation of some Western Australian work by
other laboratories. A mass spectrometry unit
established at the Department of Geology of the
University of Adelaide in 1972 has published
Rb-Sr results from the eastern Yilgarn Block
(Cooper et al. 1976), and has also worked in the
Pilbara Block. Yilgarn Block Rb-Sr results have
also come from commercial laboratories in
Adelaide (Webb and Watts 1975), some K-Ar
work on the Albany-Fraser Province has been
carried out at the University of Queensland
(Stephenson et al. 1977).

The University of Melbourne has a varied
programme of geochronological research in
Western Australia, concentrating mainly on
fission-track techniques, but including ion-probe
isotope geochronology; no results have been pub-
lished at the time of writing. The CSIRO
Division of Mineral Physics also has current
plans for both Rb-Sr and Sm-Nd studies on
Western Australian material.

26

Journal of the Royal Society of Western Australia, Vol. 62, Parts 1-4, 1979.

Figure 1. — Chronological summary up to 1977 of all publications including first reports of ages of Western Aus-
tralian Precambrian rocks and minerals by methods based on radioactive decay. Selected second reports
are also included where these augment or upgrade the first.

Thus, at the time of writing, five geochrono-
logical laboratories have continuing programmes
in Western Australian, while other work is being
carried out from time to time. Current planning
has the advantage of a reasonably well estab-
lished time-scale of years for the evolution of
the Precambrian rocks.

Publication summary

We have elsewhere presented a complete listing
of all first, and some subsequent, publications
reporting the results of geochronological work
on the Western Australian Precambrian
(Trendall and de Laeter 1978). Figure 1 is a
graphical summary of the 77 such papers there
included. It reflects clearly the phased develop-
ment of geochronological work described above.

Interaction with geological work

This brief account of the interaction of geo-
chronology with general geological studies must
accept, so far as geology is concerned, various
review papers as representative of the thinking
at the time of their appearance. The situation
shortly before the advent of geochronology was
well summarised by the then Government
Geologist, A. Gibb Maitland (1906). He assigned
all “those gneissic, granitoid, and schistose rocks,
which” . . . “form the floor upon which the
newer strata have been laid down” to the
Archaean; his next younger time division was
the Cambrian, within which were included, under
the name “Nullagine”, many of the younger,
unaltered, and undeformed sediments, now
known to have a wide range of Precambrian
ages.

Naive as this view may now seem, a modified
form of it persisted until its final concurrent
destruction by mapping and geochronology in
the 1960s. The modification consisted in accept-
ing a two-fold time division of the Precambrian
into an older Archaeozoic (Archaean) and a
younger Proterozoic; rocks of the two ages were

normally separated by a major unconformity.
This modified time classification was used, for
example, by Maitland (1919) in a subsequent
summary of the geology; in this summary
Maitland made no reference to Simpson’s (1911,
1912) published attempt to determine the age of
uranium-bearing minerals.

In 1930 E. de C. Clarke reviewed the Precam-
brian geology of the State in a Presidential
Address which was published in the following
year (Clarke 1931) and illustrates well certain
points we wish to emphasize here.

Firstly, Clarke (1931, Table p. 187) essentially
accepted the two-fold division of the Precam-
brian, with a “Great unconformity” dividing
various older rocks below from the “Nullagine
Series” above. Secondly, the Nullagine Series
was correlated between widely separated areas
of the State with no Precambrian outcrop
continuity — specifically between the Meeka-
tharra, Kimberley, and Warburton areas, and
by implication the Nullagine district itself.

Thirdly, although Clarke noted that “there is
no certainty in correlating similar rocks of the
different districts, many of which are widely
separated”, he also stated that “Correlation”
. “can only be made after much fresh petro-
logical detail combined with field observations
is available”. Like Maitland, he did not refer
to Simpson’s results, nor did he give any indica-
tion either of being aware of Cotton’s (1926)
or Holmes’ (1927) calculated ages from them,
or of appreciating the potential value of age
determinations based on radioactive decay.

These three points, and among them we
emphasize the third, apply to all general reviews
of the Precambrian geology of Western Australia
published during the 20 years following Clarke’s
(for example: Forman 1937, Clarke 1938

McKinstry 1945, Prider 1948). Thus Holmes’
publication in 1927 of the Wodgina age, on the
basis of Simpson’s 1910 analyses was met within
Western Australia by a total lack of published
reaction.

27

Journal of the Royal Society of Western Australia, Vol. 62, Parts 1-4, 1979.

The first published reference to the important
potential of geochronology for Precambrian
correlation in Western Australia was made by
Prider (1952), in a summary of the geological
structure of the southwestern part of the State.
However, two more years passed before Prider
(1954) made the first published reference within
Western Australia to Holmes’ (1927) calculation
of an age from Simpson’s analyses, in a review
that summarized all available geochronological
information to that date.

In the absence of locally published comment
between 1927 and 1952 it is not easy to establish
whether Holmes’ publication of the Wodgina age
was in Western Australia largely overlooked, or
whether it was generally known, but not taken
very seriously. For information on this point
we are indebted to Mr. F. G. Forman. Forman
(pers. comm. 1978) is emphatic that Holmes’
work was well known to geologists working on
the Precambrian at that time, but that there was
widely and deeply felt mistrust of the reliability
of radiometric age determinations. Prider has
pointed out to us that an unpublished handbook
to accompany historical geology lectures at the
University of Western Australia in 1939 referred
to an age of 1 260 million years from a pegma-
tite in the North-West, although no authority
was cited. We accept that the lack of published
references to Holmes’ (1927) work was not due
to lack of either knowledge or interest, but to a
doubt shared by most geologists as to its validity.
The same feeling is probably echoed for
Australia generally by David and Browne (1950),
who mentioned the Wodgina age, without
citation, and with the unelaborated comment
(p. 4) “the results are not regarded as reliable”.

After Prider ’s (1954) first listing of available
geochronological results, quickly followed by that
of Wilson (1958), the application of geochrono-
logical results became common among Precam-
brian geologists in Western Australia. However,
the earlier mistrust in them took some time to
diminish, and although individual geologists,
notable among them A. F. Wilson, enthusiasti-
cally advocated their utility, general acceptance
by the majority of geologists working on the
Precambrian was finally achieved only when the
results were evidently consistent with the time
relationships demonstrable from traditional
kinds of geological evidence. The best such
example is the demonstration by Geological
Survey regional mapping during the early 1960s
(Halligan and Daniels 1964) that two sedimen-
tary successions, traditionally correlated as
“Nullagine”, were separated by a major regional
unconformity related to intense folding and
metamorphism of the older; later geochronology
(Compston and Arriens 1968) confirmed this
time relationship. Elsewhere within Australia a
similar demonstration of the concordance of
geological and geochronological evidence was
given by McDougall et al. (1965).

Comment, discussion, and the future

From the sequence of events we have traced
above, a picture emerges for Western Australia
of the reluctant acceptance of physical geo-

chronology by a sceptical geological community,
as a result of the initiative and enthusiasm of
physicists and chemists. Simpson’s attempt in
1910 to obtain a U-He age for “pilbarite” was
in the vanguard of contemporary international
effort, but was not locally appreciated as such.
The discovery by Cotton and Holmes, after an
interval of more than 15 years, that Simpson’s
published analyses could be used to calculate
chemical lead ages, met a similarly cold response.

We initially found it hard to account for this
indifference by geologists, especially during the
quarter of a century between 1927 and 1952, to
results that, judged on their presently known
merits, might be assumed to have been challeng-
ing and exciting. For a possible explanation
we are indebted to Burchfield (1975), who has
traced, in an international context, the bitter
dispute concerning the age of the earth that
continued through the latter part of the nine-
teenth century between physics, in the person of
Lord Kelvin, and geology. Kelvin, from thermo-
dynamic arguments, proposed a severely
restricted age, which geologists, from a variety
of uniformitarian considerations, could not
accept.

The discovery of radioactivity undermined
Kelvin’s arguments, and discredited, in the eyes
of geologists, the reliability of theoretical
physical reasoning as applied to the age of the
earth. It was ironic that, within a decade of
its discovery, this very phenomenon, that had
finally justified half a century of spirited geo-
logical opposition, became the basis for physical
estimates of the age of the earth which were
now unacceptably long, rather than too short.
Unhappy to embark on a further half-century
of debate, “most geologists simply ignored the
new physical results” (Burchfield 1975, p. 179).
We accept the apparent indifference of most
Western Australian geologists to geochronology
as a microcosm of this international situation;
it was in essence a negative reaction to the
earlier failure of physics, as perceived by
geologists, to make a helpful contribution to
the problem of the age of the earth.

It is all the more remarkable, in view of this
indifference, that the geochronology group
established at the Physics Department of the
University of Western Australia, was so out-
standingly successful, in any terms in which
scientific attainment is measurable; its achieve-
ments are one of the highlights in the history
we have traced in this paper. From discussion
with many of those concerned, we conclude that
its success cannot be ascribed to any single
factor, but that a fortuitous coincidence of many
unrelated circumstances combined to produce the
final result. Apart from the chance visit of
Oliphant, it was fortunate that his suggestion
was taken up by a leader, Jeffery, whose infec-
tious enthusiasm was vital to attract and sustain
the interest of able and dedicated students such
as Compston, Greenhalgh, and Riley. The prior
existence of instrumental skill within the
Department was a further prerequisite for
success, as was the availability of emergency
finance in 1953 when the project was all but
abandoned. Finally, it was significant that

28

Journal of the Royal Society of Western Australia, Vol. 62, Parts 1-4, 1979.

Precambrian granitic rocks with an age of about
2 600 m.y. which had been affected by a thermal
overprint at about 600 m.y. were available for
study. It was the challenge of this situation
that produced the Rb-Sr total rock method,
which in turn established the laboratory as an
international leader in geochronology.

Burchfield (1975, p. 216) has commented that
“From the time when Kelvin coupled the first
announcement of his results to an attack upon
uniformitarianism, until the final acceptance of
radioactive dating nearly seventy years later,
the question of the earth’s age was never
entirely free from some degree of tension
between physics and geology”. Our examination
of the Western Australian situation suggests
that, although there were earlier exceptions to
which reference has been made, the complete
reconciliation of geology and physics in this
area did not come until the 1960s. The com-
plete fusion of geological and physical skills is
now widely accepted, and unreservedly so by us,
to be a prerequisite for effective research in
geochronology, and it is interesting that this is
true both in current practical experience, and in
the context of history.

It is inevitable that a historical examination
of the kind we have attempted here should lead
us to look at the future. There are at least 16
primary natural radioactive nuclides (Rankama
1963), 5 of which have been extensively used in
Precambrian geochronology: 40 K, 87 Rb, 232 Th,
235 U and 238 U. In Western Australia, the 87 Rb- 87 Sr
decay system has provided the main basis for
establishing the broad time framework which is
now known with some confidence. There is
much current interest in U-Pb and Th-Pb
methods, and the application of these, particu-
larly to zircons and other minerals, is likely to
increase. Among the remaining possibilities
187 Re and 176 Lu have been partially developed as
geochronometers (Hirt et al. 1963, Von Herr et al.
1958) , but have not yet been applied to Western
Australian samples. The decay of 1,7 Sm to 143 Nd
is currently emerging as a decay scheme with the
special advantage of “seeing through” severe
geological disturbance of a sample (McCulloch
and Wasserburg 1978), and it is likely that this
method will be applied to the Western Australian
Precambrian in the near future.

The radioactive clocks, which are the key to
our understanding of the chronology of the earth,
have progressively yielded their secrets as tech-
nological developments have enabled minute
quantities of the radioactive elements and their
daughter products to be measured with ever
greater accuracy and precision. There seems to
be no reason to doubt that further refinements
are possible as an ever-increasing number of
chronometers are available singly or in combina-
tion to tackle the problems of Precambrian
geology.

Acknowledgements . — Thanks are extended to the
following persons who provided us with relevant infor-
mation or comment during the preparation of this
paper: Prof. G. M. Brown, Dr W. Compston, Dr R. M.
Farquhar, Mr K. J. Finucane, Mr F. G. Forman. Dr H.
L. James, Dr P. M. Jeffery, Emeritus Prof. R. T. Prider,

Dr G. H. Riley, Prof. J. T. Wilson, and the Director
of the Government Chemical Laboratories, Perth.
However, we remain responsible for all opinions
expressed in the paper, as well as for any errors. One
of us (AFT) has the approval of the Director, Geological
Survey of Western Australia, to publish. We finally
wish to thank Dr A. E. Cockbain, Honorary Editor of
this Journal, for convincing us that this paper would
be worth the writing, and thereby revealing to us our
own initial ignorance in an area where we had assumed
ourselves to be well informed.

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31

Journal of the Royal Society of Western Australia, Vol. 62, Parts 1-4, 1979, p. 33-43.

Aspects of Science in Western Australia 1829-1979.

A volume to commemorate the 150th Anniversary of the founding of Western Australia.

History of zoology in Western Australia

by D. L. Serventy

27 Everett Street, Nedlands, W.A. 6009

Abstract

Previous histories of zoology in Western Australia are cited (including those of
Alexander, Glauert and Whittell) and the survey continued to the present day.
Several periods of activity may be recognised, beginning with the anecdotal records
left by the early, mainly Dutch, voyagers, followed by the more substantial ones of
the professional naturalists of the English and French marine-exploring expeditions
from the late 18th Century onwards. The first colonists in 1829 included many
talented observers and collectors, largely amateur, but the next generation settlers
were so occupied with subsistence problems that intellectual life ebbed, not to be
revived until the late 1880s. Visiting and resident naturalists, and the establishment
of the State Museum, resulted in a renewed flowering of zoological activity, often by
interested laymen, and supported by private patrons. After World War I these
were superseded by an increasing number of academically-trained zoologists,
professionally employed and centred on the University of Western Australia and
the Western Australian Museum. They added ecology and physiology to the classical-
style studies in taxonomy and comparative anatomy and revived field surveys for
the documentation of the fauna of the State.

Introduction

A detailed history of the progress of zoological
discovery in Western Australia was prepared by
W. B. Alexander (1914, 1916, 1916a, 1916b, 1917),
the Cambridge zoologist, who was on the staff of
the Western Australian Museum between 1912
and 1920. His meticulous studies summarised
chronologically all publications dealing with
events between 1618 and 1840, with some
reference to an apocryphal account dating back
to 1499, accepted by some historians but
discounted by Alexander.

Major H. M. Whittell and I in our Birds of
Western Australia (in its five revised editions
between 1948 and 1976) expanded Alexander’s
treatment to modern times but restricting the
coverage to ornithological discovery. In addition
we divided the chronological account into three
fairly well defined periods: (1) the period of the
early travellers, 1618 to 1700, when laymen’s
accounts and anecdotal records predominated,
mostly of little scientific merit; (2) the period of
the great exploring expeditions, 1791-1826, when
discoveries of considerable substance were made;
and finally (3) the period of the settlement of
the colony, from 1829 onwards. This last period
was subdivided into four sections — that dealing
with the pioneer observers, 1829 to 1839; that of
the visit of John Gilbert and his contemporaries,
1839 to 1843; the so-called “barren years”,
between 1843 and 1887, and the modern period
from 1887 onward.

In 1968 I brought Australian ornithological
discovery as a whole into this framework
(Serventy 1972) but pointed out that a “barren

period” hardly existed as such in south-eastern
Australia. Intellectual life in Western Australia
ebbed rather low after the comparatively brilliant
era of the first colonists. This cultural decline,
after the passing of the first generation of
settlers — many of them well-educated gentry
carrying on the interests of their class in
England at the time — has been noted by histor-
ians in other disciplines. Thus Kornweibel
(1973) in his history of music in Western
Australia, refers to the “Doldrums” of the mid-
century years and that the revival of musical
appreciation did not really come into its own
until the 1880’s.

Various partial histories of the development
of knowledge of the Australian fauna have
appeared. One of the most substantial supple-
ments to Alexander’s general survey is that by
Glauert (1950) on the marsupials of the State.
He elaborated on Alexander’s treatment, added
omitted data and brought the history forward
to the mid-1940’s. Hedley (1916) summarised
the growth of knowledge of the Mollusca of the
State from 1801 to 1912. Australia-wide com-
pilations, with marginal relevance to Western
Australia, include Gilbert Whitley’s summaries
of researches (Whitley 1970, 1975). Important
historical data are to be found, also, in special-
ised bibliographies, including that of Musgrave
(1932) on Australian entomological writings
between 1775 and 1930. Of particular value in
this connection is Whittell’s Literature of Aus-
tralian Birds (1954) which, besides being an
index of published information up to the year
1950, contains an historical account to 1850
Though it is essentially an ornithological history
there are references to other animal groups.

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Journal of the Royal Society of Western Australia, Vol. 62, Parts 1-4, 1979.

In the present article only scanty references
will be made to works already dealt with in
Alexander’s thorough surveys to 1840, but the
few additions to his work, as the result of later
discoveries of relevant literature, will be in-
cluded. As this account is a history, not a com-
plete index to publications on Western
Australian zoology, minor or sporadic papers will
not be referred to, unless they report something
notable.

The period of the early voyages (1618-1728j

The published accounts of natural history
observations during this period, mainly by Dutch
voyagers between Holland and the East Indies,
are more of curiosity value than of scientific
worth. Alexander has listed all that are of any
relevance to this State.

The ship’s company of the Batavia, wrecked at
the Abrolhos in 1629, were the first Europeans
to study an Australian marsupial. The Tammar
at the Wallaby Islands was reported on by
Francisco Pelsaert. Some problems regarding
the ornithology of these islands were resolved
when Henrietta Drake-Brockman (1963) deduced
that the Batavia was not wrecked on Pelsart
Island as previously thought but in the Wallaby
Group further north. Ornithologists had been
puzzled why Pelsaert had not mentioned the
enormous flocks of nesting terns and other sea-
birds on Pelsart Island. Actually he never
visited it. Important new information on the
zoology of the Southern Group of the Abrolhos
Islands was found in the journal of Adriaen
van der Graeff, the second mate of the Zeewyck
which was wrecked on Pelsart Island in 1727.
He was the first to indicate that the Wedge-
tailed Shearwaters, on which the castaways
subsisted, were migratory, first returning to the
nesting islands during the third week in August
(his actual first date of their arrival being
August 20). This detail of the species’ life
history was not appreciated by modern orni-
thologists until our own times and it was
generally believed that the birds were sedentary
(Rogers 1975).

The first European to write on the Rottnest
Quokka was Samuel Volckersen, captain of the
Waeckende Boey, during his search in 1658 for
survivors of Be Vergulde Draeck, wrecked in
1656. The log of his first officer, Abraham
Leeman, has been reviewed recently by Tyler
and Ross (1977) who interpret the “gulls” on the
islet at Sand Knoll Ledge, north of Cervantes,
which Leeman’s party ate, as fledgling Wedge-
tailed Shearwaters.

Alexander (1914) has pointed out that the
English translations of the probable meanings
of birds names in the old Dutch documents, as
given in standard works such as those by Heeres
(1890) and Major (1859) “are frequently wrong,
as is only natural, since neither of these authors
were ornithologists”. Alexander sought the
advice of a Dutch naturalist, F. E. Blaauw, in
correcting them and also gave renderings for
the first time, of some Dutch names which
Heeres and Major had left untranslated.

The period of the great marine exploring
expeditions (1791-1826)

The series of great oceanic expeditions sent
out principally by the governments of Great
Britain and France began an era of scientific
exploration which added notably to the accumu-
lation of zoological knowledge of the Australian
region. Captain James Cook and Sir Joseph
Banks are the most celebrated individual figures
characterising this period but they did not
themselves visit the waters of Western Australia.
However Banks’ journal of his impressions of
Batavia (now Jakarta) in Java gives an impor-
tant contemporary view, scientific and social, of
the situation in our near neighbour to the north
(Beaglehole 1962).

The earliest of these expeditions to visit
Western Australian seas was that in the vessels
Discovery and Chatham in 1791. George Van-
couver was in command and the naturalist
was the eminent Archibald Menzies. The most
productive was undoubtedly the French expedi-
tion in the Naturaliste and Geographe, 1801-
1803, under the command of Nicolas Baudin.
Alexander (1916) had an unbounded admiration
of the ability and dedication of the principal
naturalist, Francois Peron (“as showing the zeal
and energy of this great man . . . ”) but a
rather different picture of Peron is painted by
his commander Baudin in the recently published
translation of the latter’s log (Cornell 1974).
Nevertheless there can remain no doubt of the
competence and excellent scientific work of
Peron and his associates.

The work of the several later expeditions
surveying Western Australian waters, led by
Flinders (1801), King (1818), de Freycinet
(1818), Dumont d’Urville (1826) and Wickham
and Stokes (1840 in the Beagle ) has been
reviewed by Alexander. All contributed to
knowledge of marine and island zoology.

The last of the surveying vessels operating on
the western coast was H.M.S. Herald (Captain
Sir Henry Mangles Denham) which surveyed
Shark Bay in 1858. The surgeon, Frederick
Matthew Rayner, made important collections of
bird specimens here and further north. Chrono-
logically this expedition belongs to a later
section in this history. But it may more
properly take its place here. Denham is better
known for his surveys in New Guinea and
eastern Australian waters (Ingleton 1944).

The period of the pioneer settlers (1829-1842)

Both at the Swan River, and earlier at Port
Jackson, the first arrivals among the settlers
and Government officials exhibited a remarkable
zest for natural history inquiry. Many had
shared the vogue for natural history prevalent
in England since the publication of Gilbert
White’s Selborne in 1789. A new fauna and
flora excited latent tastes in such studies. In
the east Government officials in the First Fleet
were in the forefront in documenting the natural
phenomena around them. The first book pub-
lished about Australia after the settlement,
Surgeon-General John White’s Journal of a
Voyage to New South Wales (1789), dealt
extensively with the local fauna and flora,

34

Journal of the Royal Society of Western Australia, Vol. 62, Parts 1-4, 1979.

illustrated by many hand-coloured copper
engravings. Some of the Establishment officials
became more interested in their natural history
pursuits than in their official duties. This
seemed to be the case with Colonel William
Paterson, commander of the New South Corps
when Governor Bligh was deposed: he was very
greatly attached to botanical studies — and so
highly regarded in this field as to be elected an
F.R.S.

At the Swan River, as Alexander’s literature
review indicates, there was a correspondingly
keen interest in natural things. Lindley’s A
sketch of the vegetation of the Swan River
Colony, published within 10 years of the found-
ing of the settlement, is a graphic illustration
of how thoroughly the first settlers had set about
botanical studies, largely stimulated, of course,
by the presence among them of James
Drummond. There is no similar published codi-
fication of zoological knowledge but it would be
possible to compile one.

There was keen competition among some of
the colonists for the natural history spoils
available for the inquisitive and acquisitive.
Drummond wrote rather petulantly to Sir Joseph
Hooker at Kew that a visiting German naturalist,
J. A. L. Preiss, who arrived at the Swan River
in 1838 and stayed until 1842, was buying up
natural history material from the local settlers
which Drummond felt should be his prerogative.
Preiss was an unusually productive collector :
when he left for London he had a collection of
200 000 plants over 600 bird specimens and a
big collection of mammals, reptiles, insects and
other fauna. He had offered the collection
earlier, in October 1838, to the Colonial Govern-
ment for $6 000, $4 000 in “ready money” and
the balance in the form of a grant of land,
but apparently nothing came of the offer
(Glauert 1948). The collections were dispersed
over several European museums and were worked
over by specialists in ornithology, mollusca and
botany, but the whereabouts of much of the
material is still unknown (Meise 1951). One
interesting specimen probably collected by, or for
Preiss, escaped proper scientific study until 1907
when Siebenrock, at the Vienna Museum
described what was to become known as the
celebrated Short-necked Tortoise, Pseudemydura
umbrina (Glauert 1955). It had been mis-
identified by an earlier research worker and the
assumption was that it had been part of an
original Preiss collection. The species is now
confined to two seasonal swamps at Bullsbrook.
However, it would appear that at the time of
the original Swan River settlement it had a
more extensive range. The Sydney botanist
Charles Fraser, who accompanied James Stirling
in his 1827 reconnaissance of the Swan River,
stated when at Upper Swan: “The ridges of the
banks are perforated with immense numbers of
deep pits, the origin or cause of which we could
not at first ascertain. They proved to be made
by the natives for the purpose of catching land
tortoises, with which these ridges abound” (Hay
1906). This must refer to the Short-necked
Tortoise, then aestivating underground in the
vicinity of its winter swamps.

The most significant naturalist at the Swan
River at this period was John Gilbert, an able
collector sent out by John Gould then working
on his monumental multi-volume work on the
Birds of Australia. Gilbert laid the foundations
of ornithological knowledge of the south-west
of the State. He paid two extended visits: from
March 1839 to January 1840 and from July 1842
to September 1843. Few new full species of
birds have been found since his time and he was
the discoverer of such distinctive new forms as
the Noisy Scrub-bird, Western Bristle-bird and
Western Whipbird, and provided important data
bearing on the contraction of their ranges into
the “inner” South-West since European settle-
ment. Gilbert also collected marsupials for
Gould. He is probably the only naturalist to
have seen alive in its habitat the Broad-faced
Potoroo, Potorous platyops, and he collected the
type specimens of almost half of the species and
subspecies of recent marsupials occurring in the
south-west. Many of his field notes were used,
wholly or in part, by Gould in his Mammals of
Australia, often without acknowledgement.
Gilbert’s original notes, in letters to Gould and
some notebooks, have been subsequently re-
located at odd times and in various repositories
and are still turning up (Chisholm 1964). The
latest find was in a batch of papers found in the
British Museum of Natural History, South
Kensington by Ederic Slater of the CSIRO in
1973. His status as a mammal field worker is
discussed by Whittell (1954). He joined
Leichhardt’s expedition in 1844 to Port Essing-
ton, to further his collection of natural history
material. For his abilities he was promoted in
the field to second in command but was killed
by Aborigines on June 28, 1845.

Lieut. George Grey, later knighted during a
distinguished career, as governor and statesman,
in other States, New Zealand and South Africa,
explored extensively in Western Australia
between 1836 and 1840, collected actively for
overseas specialists, and published his observa-
tions in two volumes (1841) which are a valuable
account of zoological knowledge of the Colony
in that period. Observations on the fauna are
included in a chapter on the food and hunting
of the Aborigines, and there is a catalogue of
birds, reptiles and amphibia, with an account
of some insects from King George Sound. There
were eight plates of reptiles and amphibia and
11 figures of insects. Grey’s book is reviewed
by Alexander.

A wealth of natural history information is
contained in the publications of George Fletcher
Moore (1832, 1842, 1884 and various articles in
the Perth Gazette). Moore was a substantial
colonist from Ireland who emigrated in 1830 in
the hope of obtaining an official legal appoint-
ment as well as becoming a landed proprietor.
Before he returned to England in 1852 he had
achieved both objectives— he had become the
first judge in a civil court, Advocate-General
a member of both the Executive and Legislative
Councils, and was acting Colonial Secretary on
the death of Peter Broun, as well as being the
owner of about 10 000 ha of land in fee simple

Journal of the Royal Society of Western Australia, Vol. 62, Parts 1-4, 1979.

and several allotments in towns. His writings
reveal him to be an unusually acute field
naturalist.

Moore’s Descriptive Vocabulary (1842) of the
local Aborigines is not only a glossary of many
terms in use in various places in the South-West,
but so annotated as to provide natural history,
anthropological and geological information in a
succinct way. It is not wholly Moore’s work.
The manuscript had been placed in his hands
“in a very crude state by Governor Hutt, that
I should get it published in England.” The data
in it had been collected by Francis Armstrong,
Sir George Grey, Charles Symmons, the Bussell
brothers from the Vasse and an anonymous
person who “assumed the name Lyon.” After
he had edited and enlarged the manuscript
Moore commented that “I might fairly and truly
use the familiar expression, ‘Quorum pars magna
fui’.”

The letter-diaries (published in 1832 and 1884)
contain more extended natural history informa-
tion, which has been reviewed by W. B.
Alexander. Information in some missing letters
(e.g. in those of the latter half of 1833), and
hence unavailable to Alexander, is contained, in
part, in the Perth Gazette of the period. In the
issue of February 1, 1834, in a report by Moore
of an excursion which determined the connection
of the Avon with the Swan River, there is
botanical but no zoological information.

Moore was with Ensign Robert Dale when they
collected the first specimen of the Numbat near
Mt. Kokeby in September 1831.

The barren years (1843-C.1880)

With the passing of the first generation
settlers and the activities of their descendants
being almost wholly absorbed with economic
survival there was comparatively little leisure or
energy left to be devoted to such pursuits as
natural history. However, some noteworthy
projects were initiated by officers of the
Imperial service and other visitors.

In January 1852 William Ayshford Sanford
took up appointment as Colonial Secretary and
remained until 1855. A Cambridge man, lie was
interested in ornithology and other zoological
studies, becoming after his return to England a
well-known palaeontologist, working with Pro-
fessor Sir William Boyd Dawkins on the British
Pleistocene Mammalia. In Perth his most
noteworthy achievement, in zoology, was the
organization of the Austin surveying expedition
to the Murchison in 1854 and working up its
natural history results (Austin 1855). This
expedition was instrumental in the discovery of
the Night Parrot near Mt. Farmer (Serventy
1968). Also Austin’s journal shows that they
saw the Ghost Bat Macroderma gigas at Mt.
Kenneth, though this fact was not recognized
until years later by Glauert (1950). This would
be the first observation of the species in Aus-
tralia, about a quarter of a century before
Dobson formally described it from Queensland.

Another visitor to the State who published on
the natural history of the country at this time
was Bishop Rosendo Salvado, of New Norcia. His

book on Western Australia, with chapters on the
flora and fauna, was published in Rome in 1851,
five years after he had arrived in Western
Australia. Its first English translation did not
appear until 1977 (Stormon 1977). The zoo-
logical matter in the book is not of very great
importance but some interesting direct personal
observations are contained in it, intermingled
with information garnered from other published
accounts, not relevant to Western Australia.
There are citations of names and affinities which
are not always accurate. Present-day workers
are puzzled at the origin of some of the scientific
names used by Salvado. Some originate with
George Fletcher Moore, 1842 — Anas novae-
hollandiae for the Black Duck and Formica
maxima for the “Lion Ant” or Bullant. But
Pelecanus spectabilis is a “new” name for the
Pelican. It is still unknown how either Moore
or Salvado came by these names. Father
Stormon suggests that “spectabilis” might be a
slip for “ conspicillatus” , the correct specific
name for the Pelican.

Among the more valuable personal observa-
tions by Salvado is one worth repeating in full:
“Not seldom in the winter season, that is, June
and July, I have found birds big as a thrush,
which had fallen to the ground benumbed by
the bitter cold of the night; I would take them
up in my hands and not let them go until they
had recovered their natural warmth.” Salvado
has here clearly described a phenomenon, of
over-night torpidity, which has only very
recently come under the attention of Australian
ornithologists (Serventy 1970, Ives 1973). The
phenomenon must, however, be of very general
occurrence, and will no doubt be the subject of
many specialist studies in the future.

This generally bleak period of zoological
enterprise was brightened somewhat in 1866 and
again in 1868-69, when George Masters was sent
by the Australian Museum, Sydney, on collecting
trips to the south coast. He obtained several
specimens of the now dwindling Noisy Scrub-
bird and a large collection of mammals (listed
by Glauert 1950).

This period may be said to merge with the
next, the Modern Period, with the era of William
Webb, of Albany. He worked in isolation from
the main communities in Western Australia, but
his collecting of animals and plants brought
him into contact with eastern states specialists,
including Baron von Mueller and W. S. Macleay,
who had founded the Macleay Museum in the
University of Sydney. Webb also sold natural
history material to passengers on overseas ships
calling at Albany. The well-known botanical-
artist traveller, Marianne North, met him in
1880 and provided the information that Webb
had been a convict and was married to an
Aboriginal wife. After The Albany Mail began
publication in 1883 Webb contributed a series of
useful natural history articles, the ornithological
ones of which are summarised by Whittell, who
cited others by title (Whittell 1954).

36

Journal of the Royal Society of Western Australia, Vol. 62, Parts 1-4, 1979.

The modern period (1887 onward)

Introduction

After the English ornithologist Thomas Carter
arrived in Western Australia in 1887 there began
an era of continuing activity in ornithological
studies in the State and there was an accom-
panying crescendo of general zoological activity.
Carter, a pastoralist by calling, was a talented
amateur. However, he worked with professional
ornithologists both in England and Australia.
The important thing was that there was now
to be a continuum of contact between ornitho-
logists to the present day.

In addition a number of expeditions, from
overseas and within Australia, were mounted to
explore the natural history of the inland areas
and some islands.

More organized scientific activity became
possible when W. B. Alexander joined the scienti-
fic staff of the Western Australian Museum in
1912 and the University of Western Australia
was founded in 1913, with Professor W. J.
Dakin in the chair of biology. Later still, with
the re-vitalising of the University Zoology
Department by the arrival of Professor H. W.
Waring in 1948, the building up of the CSIRO
Divisions of Fisheries, Wildlife and Entomology,
the appointment of Dr. W. D. L. Ride to the
directorship of the Western Australian Museum
and the setting up of research staffs in the
Fisheries and Fauna (later Wildlife) Depart-
ment, there was a spectacular upward spurt in
the quantity and quality of zoological research.
This is continuing with the increase in number
of tertiary education institutions.

The Modern Period may be subdivided as
follows: (1) the building-up era, to the first
World War; (2) under the leadership of the
University and Museum, from about 1912 to
about 1950; and (3) the present-day era of
sophisticated zoological research.

From the arrival of Thomas Carter to the
eve of World War I, 1887-1912

Carter did extensive field work in the region
of North West Cape, where his sheep station
(Point Cloates) was situated (Vines 1968). He
laid the foundations of ornithological knowledge
of the North-West and early recognized the
existence of opportunistic breeding dependent on
rainfall. His work overlapped that of another
English migrant to the State, who, like him, had
had previous experience in modern ornithological
practice at home. This was F. B. Lawson
Whitlock whose collecting on behalf of various
patrons extended knowledge of bird distribution
and his material provided the basis for taxo-
nomic work on subspeciation by Gregory
Mathews in England. A Victorian ornithologist,
A. J. Campbell, visited the southern part of
the State in 1889 and a series of reports by him
filled in important details of distribution at that
time (Campbell 1890).

Two inland expeditions, organized from beyond
the State, collected zoological material in the
inland areas. The Elder Expedition of 1891 had
as naturalist Richard Helms, a remarkably able
and versatile German migrant whose collecting-
in several animal groups was the “chief source

of knowledge as to the fauna of the dry interior
regions of Western Australia” (Alexander 1916).
Helms later worked in Western Australia and
between 1896 and 1900 was biologist to the
Department of Agriculture. His publications on
the East Kimberley and the Abrolhos Islands
are particularly important. The other expedi-
tion, the Calvert Expedition of 1896 with G. A.
Keartland as naturalist, was concerned mainly
with ornithology.

Two expeditions from Scandinavia worked in
the north of the State. Knut Dahl, a Norwegian,
collected in Dampierland in 1895 and added to
the knowledge of local birds and mammals
(Glauert 1950, Dahl 1926). The Swedish
Scientific Expedition of 1910-13, under E.
Mjoberg, also worked in the Kimberley Division
but as well visited the area between Perth and
Albany. Its results were published in 32 parts of
the Svenska V etenskapsakademiens Handlingar
between 1913 and 1922. The expedition ornitho-
logist, R. Soderberg (1918) was the first to draw
attention to a peculiar phenomenon of the arid
areas of Australia — the simultaneous occurrence
of moult and breeding — an adaptation con-
sequent on irregular opportunistic breeding.

Two considerable British expeditions and one
minor one visited the State in the early years of
the present century. The era was dominated, to
some extent, by Oldfied Thomas, the mammalo-
gist of the British Museum, who was instrumen-
tal in setting up the Balston Expedition of the
British Museum in 1904-1905 with G. C. Short-
ridge as collector. He worked in the southern
part of the State, on birds and mammals, mak-
ing substantial contributions to distributional
knowledge (Ogilvie-Grant 1909, Shortridge 1909,
Glauert 1934, 1950).

In 1912 P. D. Montague, a Cambridge zoologist,
visited the Montebello Islands (Montague 1913).
The general collections were worked on by
specialists and published in the Proceedings of
the Zoological Society of London in 1914 — Hogg
(spiders), Montague (birds, reptiles, fishes and
some insects), Rathbun (Crustacea), Iredale
(molluscs) and Robson (cephalopods) . Montague
lost his life in World War I.

When Radcliffe-Brown, from Cambridge,
visited the State in 1910 on an anthropological
expedition, he was accompanied by a young
Cambridge graduate, E. L. Grant Watson, as
zoologist. Watson collected mainly insects and
his account of the enterprise (Watson 1968)
would interest present-day expedition members
who work under grants or salaries with generous
daily field allowances. Watson received no
salary and was expected to pay his way by
selling specimens he obtained to specialists. He
had made arrangements, for example, to receive
ten shillings for 20 specimens of beetles, more
for any considered rare, and a pound for 12 fleas
of any species!

The most productive expedition of this period
was the Hamburg South-Western Australian
Research Expedition of 1905, under Dr. W.
Michaelsen and Dr. R. Hartmeyer. With the
exception of the Swedish expedition of Dr
Mjoberg and Montague’s Montebello expedition^
this was the first fully professional scientific

Journal of the Royal Society of Western Australia, Vol. 62, Parts 1-4, 1979.

expedition of this period — in point of time actu-
ally the first. The expedition was mainly
concerned with invertebrates but collected also
reptiles, amphibia and some fishes. The results
worked on by teams of specialists appeared in
Die Fauna siidwest-Austr aliens, in 5 volumes
from 1907 onwards (Michaelsen and Hartmeyer
1907-1930). Three reports on the results and
abstracts of most of the first three volumes were
published locally by Michaelsen (1908, 1911 and
1914). The reports of this expedition provide
the first systematic studies of many important
invertebrate groups including Annelida, Crusta-
cea, Onychophora and others.

An important generalisation on the relations
of the Western Australian fauna to that of the
rest of Australia was made by Professor Baldwin
Spencer, of the University of Melbourne, after
reviewing the results of the Horn Scientific
Expedition to Central Australia (Spencer 1896).
He introduced the concept of the three faunas —
Bassian, Eyrean and Torresian — which has been
generally adopted ever since, with modifications
(cf. Serventy and Whittell 1976, Serventy 1972,
1973).

From the long-term point of view the most
significant happening in this epoch was the
establishment of the natural history section of
the Perth Museum in 1892 under the curator
Bernard H. Woodward. In 1895 its name was
changed to the Western Australian Museum
(Woodward 1903). Woodward, director of the
institution since 1890, energetically encouraged
the building-up of its zoological collections even
though he himself, primarily a geologist, was
mainly concerned with the arts (he was director
also of the Art Gallery) . In 1894 the Museum
engaged the services of a taxidermist O. H.
Lipfert, an unassuming but extremely competent
naturalist whose role has been generally under-
estimated. He trained J. T. Tunney as a field
collector who between 1895 and 1906 made
extensive collections of mainly birds and mam-
mals throughout the State. Lipfert himself also
collected widely, his last considerable expedition
being with the Canning Stock Route recondition-
ing party between 1930 and 1931 when he was
over 65. But his main occupation at the
Museum was the setting up of the displays of
mammals, birds and other exhibits in the public
galleries. One of the most outstanding was the
diorama of the animals near Perth which he
set up at the turn of the century, long before
such displays became general in overseas
museums.

Woodward sought the aid of outside natur-
alists in running specialist museum activities.
The most important was A. W. Milligan, a Perth
law clerk, who between 1901 and 1908 held the
appointment of Honorary Consulting Ornith-
ologist. He went on numerous expeditions and
described the new species of birds that came to
the Museum, the most notable of which was the
Black Grass-Wren from the Kimberley Division,
collected by Dr. F. M. House in 1901 during the
F. S. Brockman survey expedition. This bird,
though sought for assiduously in the intervening
years, was only rediscovered in 1968 by the

Harold Hall Expedition of the British Museum
(Freeman 1970). Milligan made important,
though often faulty, generalisations on bird
distribution in the southern parts of the State.

Though he was not a zoologist on the Museum
staff at this period mention should be made of
the work of Ludwig Glauert between 1909 and
1915 on the cave fossils of the South-West. He
captured the popular imagination by showing
that such creatures as the Koala, Tasmanian
Thylacine and Devil, wombats and other animals
thought to be peculiar to south-eastern Aus-
tralia, had peopled the South-West in Pleisto-
cene and Sub-Recent times. His public lectures
on the finds attracted influential audiences,
including the Governor of the day. Glauert was
transferred from the Geological Survey to the
Museum in 1910 and his zoological work will be
referred to in the next section.

Another zoological institution came into being
in this period — the South Perth Zoological
Gardens. The gardens were laid out by the first
director, Lieut-Col. E. A. Le Souef in 1897. Le
Souef was a member of the renowned zoological
gardens family of Melbourne and his brother
founded the zoological gardens in Sydney. The
history of the South Perth institution and its
zoological and acclimatisation activities has been
written by Jenkins (1978).

Finally, the Fisheries Department was con-
stituted, following a report on local fisheries
administration by Lindsay Thompson, of New
South Wales (Thompson 1898). The subsequent
reports by the Chief Inspectors (Gale 1900-1911,
Aldrich 1912-1933, Fraser 1938-1939), including
those of their regional inspectors, Abjornsson
and others, provide a wealth of data on the
condition of the estuaries and other waters which
have been undeservedly overlooked by later
research workers. The second Chief Inspector,
Fred Aldrich, who held office between 1911 and
1938, was as self-effacing a man as was Lipfert
at the Museum, but had conducted useful inves-
tigations the results of which are still mostly
concealed in departmental files. Chittleborough
has recently published one of his pieces of work,
on the marine crayfishes (Chittleborough 1967).

In addition to these departmental reports
much informative data on fisheries are contained
in the minutes of evidence of Parliamentary
inquiries into various aspects of fishing, pearling
and whaling (e.g. Baglin 1922, Daglish 1906,
Holman 1915 and Leslie 1948-1949).

In this period, too, began the first of the
State’s natural history and biological societies.
The earliest was a purely botanical group, the
Mueller Botanical Society, founded in 1897,
which in 1904 was enlarged in scope to become
the Western Australian Natural History Society.
In 1909 it was transformed again to become
the Natural History and Science Society of
Western Australia. All three bodies regularly
published their proceedings and papers. In 1913,
consequent on events which will be recounted
in the next section, the Society became the
Royal Society of Western Australia.

38

Journal of the Royal Society of Western Australia, Vol. 62, Parts 1-4, 1979.

The ascendancy of the University and the
Museum

Two events in 1912 and 1913 proved turning
points in the advancement of zoological studies
on modern lines in Western Australia. Bernard
Woodward retired from the directorship of the
Western Australian Museum in 1914. The post
was not filled but the collections were placed in
the custody of three Keepers. The Keeper of
Zoology was Mr. W. B. Alexander, the Cambridge
graduate already mentioned who had joined the
museum staff as an assistant in 1912. Alexander
vigorously expanded the collections, organized
educational displays in the museum on modern
lines and by his example stimulated modern
ornithological field studies which had a perman-
ent effect on his followers (Serventy 1967).

The other event was the appointment of
Professor W. J. Dakin to the first chair of
biology in the newly founded University of
Western Australia in 1913. Dakin was a marine
and freshwater biologist. He enthusiastically
threw himself into local researches. In pre-
liminary plankton surveys of the Swan River
estuary (Dakin 1916), he established that the
phosphorescence in the water was due to a
tintinnid ciliate protozoan, that the common
jellyfish, Aurelia aurita, produced large numbers
of ephyrae in June and that it would be possible
to work out the complete development of the
Swan River prawn from a series of plankton
catches — this last anticipation not being realised
until recent studies by Fisheries Department
biologists. In inland waters he investigated the
phyllopods of ephemeral pools and worked on
Peripatus in damp earth habitats. He organized
two scientific expeditions to the Abrolhos Islands
during 1913 and 1915, financed by the Percy
Sladen Trust. The results were published in a
series of papers in the Journal of the Linnean
Society of London (Dakin 1916a and 1919,
Alexander 1922, Fauvel 1922, Tattersall 1922,
Hickson 1922, Clark 1923, Dendy 1924,
O’Donoghue 1924). He worked in close co-
operation with Alexander of the Museum and
the two were involved with the Abrolhos
expeditions. Unfortunately both left the State
for other spheres in 1920.

It was during their period in Perth that Dakin
and Alexander were associated with the evolu-
tion of the old Natural History and Science
Society into the Royal Society of Western Aus-
tralia. This happened in 1913. Dakin became
the first president and Alexander the second
secretary, following a short spell in that office
by Mr. M. A. Browne.

After the departure of Dakin and Alexander
their posts were filled, respectively, by G. E.
Nicholls at the University, and by L. Glauert at
the Museum. They did not form the same
co-operative team as did Dakin and Alexander,
but nevertheless both continued the research
tradition laid out by their illustrious prede-
cessors. Nicholls (a London University
graduate) who held the biology chair from 1921
to 1947, was essentially a comparative anatomist
of the classical school and the graduate research
in his department was mainly taxonomic. His

own research interests were in the lower
Crustacea. He was stimulated by Glauert’s
discovery of a local species of the distinctive
freshwater isopod, Phreatoicus, and came to
specialise on the group, ultimately producing a
notable monograph on the phreatoicids from the
whole of Australia (Nicholls 1943, 1944). His
research students and other associates worked
on other crustacean groups. Earlier papers by
Sayce (1903), Wolf (1911), who worked on the
collections made by the Hamburg Expedition of
1905, and by Dakin (1914) had drawn attention
to the rich and varied fauna of phyllopods
occurring in the inland claypans and lakes.
These were collected and came under study.
Milner (1929) worked out a collection of
anostracans made by Nicholls himself; K. C.
Richardson made a study of conchostracans
which remains in manuscript in the Zoology
Department; a visiting Swedish specialist on
the anostraca, Folke Lindner, was based at the
Department in 1936 and 1937 and conducted
extended field work which contributed to a
substantial part of his massive world revision of
the order (Lindner 1941). Nicholls’ review
(1933) of the fauna of Western Australia is a
valuable pointer to the existence of a rich fauna
of damp earth and aquatic invertebrates.

Glauert’s activities at the Museum between
the two world wars were severely hampered by
reduced finances, made particularly acute by the
Depression. He had to operate virtually single-
handed and the widespread collecting which had
been a feature of Museum practice before the
first World War had to be suspended until
revived again later under a new director.
Glauert’s difficulties during the period and his
achievements in spite of them are reviewed in a
valedictory article published when he retired
from the Museum (Serventy 1957). One minor
publication of his, a Museum leaflet proposing
popular names for local mammals (Glauert
1928) attained almost instant success in popu-
larising a new name for the Rottnest Island
wallaby Setonyx brachyurus. This animal had
hitherto been referred to as the Short-tailed
Wallaby in scientific literature and “Wallaby”
by the public. Glauert selected the King George
Sound Aboriginal name, Quokka, for it, though
the Swan River native name was Ban-gap, or
Bunkup as the White settlers varied its pro-
nunciation. “Quokka” became so rapidly
accepted, through Glauert’s publicity (he was
regularly interviewed by the daily press on
current zoological news) that most people now
consider this name to have been always in use.

This period may be said to mark the with-
drawal of the amateur naturalist from his
hitherto prominent role as leader in zoological
studies in the State and his replacement, natur-
ally enough, by the professional — the trained
and better-informed staff and students of the
new University, staff members of the Museum
and of the scientific branches of State govern-
ment departments. Later still academic quali-
fications became an almost essential require-
ment for entry to the professional ranks.

39

Journal of the Royal Society of Western Australia, Vol. 62, Parts 1-4, 1979.

However, the value of the amateur did not
disappear. His role changed. In 1924 an
organization ostensibly catering for amateur
naturalists was founded in Perth, the Western
Australian Naturalists’ Club, but it was not until
nearly a quarter of a century later that it was
able to produce a periodical, the Western Aus-
tralian Naturalist, now in its 14th biennial
volume. The Club did not solely remain the
prerogative of the amateur, as popularly under-
stood, but proved to be a happy amalgam of
the activities of serious amateurs, professionals
and interested laymen, continuing the British
and European tradition of enrolling persons
finding personal pleasure in the study of the
natural sciences as well as earning their liveli-
hood in them. The German word, Liebhaberei,
aptly describes their pursuit. The history of the
Club and its achievements has been written by
Erickson (1964, 1974). The new role of the
amateur in the modern order has been discussed
by Serventy (1972, 1973).

Coincidental with these changes of emphasis
between amateur and professional in the devel-
opment of zoology in the State, which were
subtle and gradual, was the disappearance of the
private patron, or gentleman collector. In the
absence of appropriate institutions these people
had virtually subsidised natural history in the
old days, not only by sending out collectors at
their own expense as did John Gould in England
and W. S. Macleay and H. L. White in New
South Wales, but paid visiting and resident
collectors for specimens. Both patron and field
collector were dedicated to advancing natural
history knowledge and commercial profit was a
very low priority, if considered at all. The
example given previously of how a keen young
Cambridge graduate, E. L. Grant Watson, was
able to come to Australia in 1910 for a pittance
was not unique. That class of enthusiast has
by no means disappeared, but changing opinions
on collecting and the attitude of many Museums
have contributed to a situation where natural
history studies involving field collecting have
become virtually a “closed shop” to institutions
and their members.

Parallel with these changes and no doubt
related to them in a subtle way, is the altered
attitude of the public to Zoology in this State.
Mr. Jenkins in his presidential address to this
Society in 1965 (Jenkins 1965) drew attention to
the fact that membership of the Royal Society
remained virtually static between 1923 and 1965.
However in earlier years the “influence they
wielded was disproportionately large.” He
attributes this to the fact that in early years
the membership ranks included most of the
influential people of Perth — the leading states-
men, members of the judiciary, leading clergy
and merchants and socially prominent citizens.
In recent times membership is “attracting
mainly working scientists.” The annual general
meetings and conversaziones used to be impor-
tant events in Perth’s social calendar and the
Society’s ordinary meetings were reported
regularly and fully in the morning newspaper.
A presidential address by Bishop (later Arch-
bishop) Riley in 1907 was published in full and

other meetings might receive up to two columns
of newspaper space. Other scientific news
received similar treatment. Lectures at the
Museum by Glauert in 1914 on the cave fossils
of the South-West drew large audiences, which
included the Governor, Sir Gerald Strickland,
with extensive press coverage. Another lecture
by Glauert to the Western Australian Natural-
ists’ Club in 1925, introducing to the Perth
public the concept of continental drift as an
explanation of fauna distribution, occupied a
full column. Interviews with Perth zoologists,
such at Glauert and Lieut. -Col. E. A. Le Souef,
of the South Perth Zoo, on contemporary
zoological news were commonplace. In recent
years such reports became brief and occasional,
and usually restricted to bizarre happenings of
the “man bites dog” category. But latterly the
publicity tide shows signs of turning. Specialist
reporters on the West Australian like
Catherine Martin (medical biology), Alex Harris
(general science) and Michael Zekulich (agri-
cultural science) cover the local scientific scene
intelligently and in depth.

The modern sophisticated period of research

As with the beginning of the previous epoch
the present one, now in progress, coincided
approximately with the appointment of new
heads to the Zoology Department of the Uni-
versity of Western Australia (in Professor H.
W. Waring) and of the Western Australian
Museum (in Dr. W. D. L. Ride). The rejuvena-
tion of research and exploratory activity in both
institutions, made possible also by an increase
in funding and in staff numbers, has been
dramatic. This new epoch was also notable for
the increased extension into this State of the
research activities of the CSIRO (in fisheries
and oceanography, wildlife and entomology) and
the expansion of the research programmes with-
in the State Departments of Agriculture and
Fisheries and Wildlife. The progress in research
activity was accelerated when additional tertiary
educational institutions were formed — the

Western Australian Institute of Technology and
Murdoch University.

Professor Waring (Liverpool University), who
joined the University Zoology Department in
1948, introduced physiological and ecological
teaching and research into the University and
stimulated an active group concerned with
marsupial eco-physiology. This was centred on
a biological research station established in 1954
at Rottnest Island with the Quokka as the main
study animal. The project was extended to a
field station at Jandakot. Waring’s work was
widened by A. R. Main’s studies on the biology
of frogs. Research students from this training
centre now occupy important chairs and other
senior posts throughout Australia as well as
abroad.

Ride, fresh from a vigorous research school at
Oxford on vertebrate biology, expanded the
scope of the Museum. He demonstrated the
truth of the principle, which Glauert had been
unable or unwilling to grasp, that “people are
able to create funds”, if a worthwhile case can
be made out to government treasuries or private

40

Journal of the Royal Society of Western Australia, Vol. 62, Parts 1-4, 1979.

patrons. A large increase in funds became
available to the Western Australian Museum
which enabled the scientific staff to be vastly
enlarged allowing an active resumption of the
exploration work which had characterised the
Museum between 1895 and 1906.

The Fisheries and Fauna (now Wildlife)
Department became similarly rejuvenated under
another like-minded individual. This was A. J.
Fraser who came from the New South Wales
Fisheries Department to be Chief Inspector at
Perth in 1938 — his title was later changed to
Director. He was an administrative officer and
not a biologist, but he had a remarkable insight
into how such a department should develop as
an effective instrument into the research and
control of exploited natural resources. He
created the department as it now is, with a
trained research staff. In the early stages of the
process he enrolled outside scientists in a com-
mittee to plan research enterprise, a body which
has now become the W.A. Wildlife Authority
within the Department. Working in concert
with the Museum the Department has begun a
systematic zoological survey of key faunal areas
throughout the State. Publication of results are
made by both institutions on a scale never
attempted in the past.

The relevant activity in the zoological field,
within the Department of Agriculture, concerns
investigations on species considered harmful to
the farming and pastoral industries. The work
is based on a modern laboratory complex at
Forrestfield.

The major achievement of the CSIRO in
terrestrial zoology was the establishment of the
Wildlife Research Station at Helena Valley in
1966, and of which the present head is Dr.
Stephen Davies. The work organized there is
mainly ecological and ethological.

In marine zoology the CSIRO Division of
Fisheries and Oceanography has conducted local
research since 1943. Prior to that period some
marine surveys had been made by the Common-
wealth Government in the research trawler
Endeavour, under H. C. Dannevig (Dannevig
1913). His studies in the Great Australian
Bight extended into Western Australian waters.
The biological results, including taxonomic
studies on many groups, were published in six
volumes of reports between 1911 and 1933 by
the Commonwealth Department of Trade and
Customs. The CSIRO surveys in the Fisheries
Research Vessel Warreen (Captain C. E.
Pedersen) from 1947-1949 were extended later in
vessels of the Royal Australian Navy on hydro-
graphic and plankton investigations. The
Division operates in active cooperation with the
State Fisheries and Wildlife Department.

The new modes of transport now available to
the field worker on land, operating as a member
of a well-planned team of several specialists,
contrast markedly with the facilities available
to their predecessors. Whitlock in 1909 travelled
to Wiluna by a camel team; Carter, 30 years
after he first arrived in the North-West,
describes his first motor car trip in 1916. Four
wheel drive vehicles and aircraft make the most

inaccessible portions of the State available to
zoological survey teams and the impressive
modern publications of the Museum and Depart-
ment of Fisheries and Wildlife are now docu-
menting the fauna of the furthest ends of the
State.

But, curiously, the South-West corner in spite
of 150 years of settlement, still provides zoologi-
cal spoils of intense theoretical significance. In
1971 A. R. Main discovered Peripatus on the
Swan Coastal Plain south of Perth (Van der
Lande 1978) after it was universally believed
that this creature occurred only in the Darling
Range plateau. Rosen (1974) demonstrated his
belief in a lengthy and elaborate paper that a
diminutive fish found in a small brook near
Shannon River in 1959, and named by Mees
(1961) Lepidogalaxias salamandroides, was not a
primitive galaxiid but a member of a Northern
Hemisphere grouping, the esocoids, or pikes,
hitherto unknown from any southern continent
and presenting baffling questions as to origins.
Rosen’s suggested affinities have been treated
with some reserve by other zoologists. However,
whatever its relationships Lepidogalaxias must
be regarded as an extraordinarily interesting-
survival of an ancient fauna. It occurs now
in the extreme south-west corner of the State
which is remarkable for the number of animal
species of very circumscribed ranges — in fresh-
water fishes (Mees 1961), frogs (Main 1965),
and freshwater Crustacea of the jilgie-koonac
assemblages (Riek 1967). Its present depend-
able climate emphasises the point made by
Gentilli (1949) that throughout the period of
climatic changes during the Pleistocene this
has been a humid refuge. It deserves a thorough
zoological survey and in groups other than birds
and mammals which have been the preoccupa-
tion of many faunal surveys of the past. Lepi-
dogalaxias was first picked up by casual non-
professional collecting in 1959; it could so easily
have remained undetected to the present day.
No large area in this South-West corner should
be subject to any extensive habitat despolia-
tion — for developmental purposes — until

adequate biological surveys have been made.
The miscellany of information provided by
Nicholls (1933) on the extent of a damp earth
and freshwater fauna in the South-West, still
largely unworked, lends particular cogency to
his plea at the time for “an immediate and co-
ordinated investigation” into it.

References

Aldrich, F. (1912-1933). — Reports of the Chief Inspector
of Fisheries 1911 to 1933. Government

Printer, Perth (those from 1911 to 1912

printed, remainder in typescript).

Alexander, W. B. (1914). — The history of zoology in
Western Australia; discoveries in the 17th
Century. J . W. Aust. Nat. Hist. & Science
Soc. 5: 5-20.

Alexander, W. B. (1916).— Obituary : Mr. Richard Helms.

J. Roy. Soc. W. Aust. 1 : xxviii-xxix.

Alexander, W. B. (1916a). — History of zoology in Western
Australia. Part II. 1791-1829. J. Roy. Soc.
W. A.ust. 1: 3-69.

Alexander, W. B. (1916b) .—Notes on the birds met with
on the Swan River by Vlamingh in 1697.
J. Roy. Soc. W. Aust. 1 : 70-71.

41

Journal of the Royal Society of Western Australia, Vol. 62, Parts 1-4, 1979.

Alexander, W. B. (1917). — History of zoology in Western
Australia. Part 3, 1829-1840. J . & Proc. Roy.
Soc. W. Aust. 3: 1-33.

Alexander, W. B. (1922). — The vertebrate fauna of
Houtman’s Abrolhos (Abrolhos Islands),
Western Australia. J. Linn. Soc. London 34:
457-486.

Austin, R. (1855). — Journal of Assistant Surveyor R.

Austin in the interior of Western Australia.
Government Printer, Perth.

Baglin, F. A. (Chairman) (1922). — Report of the select
committee of the Legislative Council
appointed to inquire into the fishing indus-
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43

Journal of the Royal Society of Western Australia, Vol. 62, Parts 1-4, 1979, p. 45-51.

Aspects of Science in Western Australia 1829-1979.

A volume to commemorate the 150th Anniversary of the founding of Western Australia.

The Western Rock Lobster, Panulirus cygnus

by R. W. George, G. R. Morgan and B. F. Phillips

Western Australian Museum, Francis Street, Perth, W.A. 6000
Western Australian Marine Research Laboratories, West Coast Highway, Waterman, W.A. 6020
CSIRO Fisheries and Oceanography Division, Leach Street, Marmion, W.A. 6020

Abstract

The supervision of the rock lobster industry and the management controls
that have been introduced have been constantly reviewed so that Western Australia’s
multi-million dollar industry stands today as one of the few fisheries that can be
said to be properly managed. The management strategy has been determined by
co-operative decisions between State and Commonwealth fisheries institutions and
it has been based on reliable catch and effort data collected since the post-1945
expansion of the fishery, a broad understanding of the species biology after much
research work by many scientists, and the application of modern fisheries models
to assess levels of fishing effort and the possible effect on the stocks.

Introduction

The western rock lobster forms the basis of a
multi-million dollar industry which began as an
incidental fishery in the early 1900s when pots
were pulled by hand from sail-powered fishing
boats and the rock lobsters were mostly sold
direct to the housewife by the fishermen.

Little management of the fishery was needed
in those times even though, as early as 1897, a
regulation was introduced to prohibit the sale
of any rock lobster less than the legal size; the
regulation, announced in March 1897 set a mini-
mum legal weight of 8 oz. which was altered in
September of the same year to 12 oz.

The control and management of the fishery
has been the responsibility of the State Depart-
ment of Fisheries and Wildlife and the Depart-
ments’ management policy has always been based
on the best research information available.
Early research on the fundamental biology of
this species was commenced in the 1940s by the
CSIRO Fisheries Division through the efforts of
the late Dr. K. Sheard (Sheard 1962). Due to
the expansion of the State Fisheries Depart-
ment’s research and management unit and the
recent major effort on larvae by CSIRO Fisheries
and Oceanography Division, 6 scientists are
currently engaged in research on the western
rock lobster. At the conclusion of this present
review a full list of the publications relating to
research on the western rock lobster is presented.

The species ( Panulirus cygnus George 1962) is
endemic to Western Australia, occurring along
the continental shelf of the west coast from
North West Cape (21° 45'S) to Cape Leeuwin
(34° 22'S). Fishermen expanded their opera-
tions from the two early centres at Fremantle
and Geraldton and they now set their pots over
an extensive area from the shallow coastal reefs

to the edge of the shelf at 200 m. The present
flourishing fishing centres are scattered along
the west coast shelf at Kalbarri, Abrolhos
Islands, Geraldton, Dongara, Cliff Head, Jurien
Bay, Leeman, Cervantes, Lancelin, Fremantle
and Bunbury. The annual production is approxi-
mately 8 000 tonnes, 95% of which is exported
to the U.S.A. market as frozen uncooked tails.

In the mid 1960s, a serious concern was
expressed by the scientists and the fishermen
because the catches resulting from the rapid
expansion of boats, men and pots in the previous
5 years or so had not resulted in a proportionate
return to the fishermen. Thanks to the fore-
sight of early fisheries administrators, records of
catch and fishing effort had been gathered and
as a result of intensified research on these
records and a better understanding of the
biology of the rock lobster, rigorous limitations
were able to be placed on the total fishing effort
to halt the decline in catch per unit effort. This
was successfully achieved by limiting the number
of pots permitted by each boat and restricting
the total number of boats in the industry.

Annual production was maintained at a
reasonable level but more recent research has
shown that effort was again increasing due to
an increase in the number of pot lifts over the
season since fishermen were working more
efficiently and more frequently than they had
in the past. So in 1978 the length of the season
was reduced by 6 weeks to assist fishermen to
maintain a viable economic return.

Constant research into the best management
techniques has proved that a natural fishery
resource is capable of adequate management
and the Western Australian rock lobster industry
stands today as one of the very few fisheries in
the world that can be regarded as well managed.

45

Journal of the Royal Society of Western Australia, Vol. 62, Parts 1-4, 1979.

Because of this research and the results pro-
duced by the Western Australian scientists
studying the western rock lobster, Perth was
chosen in 1977 as the venue for a week-long
Australian-American “Workshop on lobster and
rock lobster ecology and physiology”, the pro-
ceedings of which have been recently published
(Phillips and Cobb 1977).

General biology and environment

In addition to the commercially important
western rock lobster, there are 6 other species
of rock lobster in Western Australia only one
of which is commercially important. This is
the southern rock lobster, Jasus novaehollandiae
that is fished in south-east Australia. In
Western Australia however, its density is not
sufficient to support more than small catches at
Esperance and Albany. Along the northern
coast, 5 species of very wide ranging tropical
rock lobsters ( Panulirus spp.) are found in
moderate concentrations in shallow reef and
coral outcrops. These species have separate
environmental preferences, are restricted in
depth to the shallow reefs and do not support
continuing catches (George 1968, 1974).

The western rock lobster occupies a subtropic
(intermediate) geographical position between
these southern and northern zones and occupies
much of the relatively wide continental shelf
that extends seaward from the west coast. The
continental shelf has been subjected to signifi-
cant sea level changes in the past and the
former coastal fringing limestone reef flats now
form lengthy submarine “ledges” that provide
shelter for the rock lobster. It is a gregarious
species and dense groups of several hundred
can be found packed closely together in the
undercut ledges of these submarine features.

The south-west swell that strikes the west
coast is a major physical factor that results in
both the constant wave action on the shallow
reefs and the deep surges that are a fairly con-
sistent feature of the submarine ledges. The
seaweed kelps provide mobile cover around and
over the entrances to the undercut ledges. The
fauna and flora associated with seagrass beds
and reef flats (molluscs, worms, coralline algae,
seagrass itself and small Crustacea) provide most
of the food supply for the rock lobsters. (Joll,
pers. comm.). Their feeding forays are usually
restricted to hours of darkness; bright moon-
light nights drastically reduce the catch in pots
and most fishermen reduce their fishing effort
during these periods. Predators on the rock
lobsters include octopus (Joll 1977) as well as
large predatory fish including sharks.

The western rock lobster is regarded as a
‘slow grower”. Chittleborough (1974d) main-
tained and grew juveniles in the laboratory,
indicating a growth-to-maturity period of about
6 years and achieved successful mating and egg
carriage under these conditions. Growth occurs
only after a moult when the old hard exoskeleton
is shed and a new soft one is expanded to its
new size by absorption of salt water. Food
requirements and the general physiology of the
rock lobster alter dramatically before, during

and after each moult (Dali 1974a, b, 1975a, b, c,
1977; Dali and Barclay 1977) and they remain
deep in their shelters until the new shell hardens
in about 7 days.

At the time of adolescence, most if not all,
individuals go through a distinct phase known
as the “white” phase. The characteristic red-
brown colour of the species is not present in the
newly moulted shell which is parchment to light
pink in colour; red pigment is gradually added
to the shell until, after about 3 weeks, they are
indistinguishable from the normal coloration.
The white phase occurs regularly in November-
December of each year in the shallower waters
and a rapid offshore migration occurs during
the “white” fishing season (George 1958b).

The sexes are separate in P. cygnus and sex
ratios in the adult stock approximate 1:1 but
because of differences in availability this ratio
may vary significantly in pot catches (Morgan
and Barker 1974). The size at first maturity of
females and the mean size of breeding females
varies significantly throughout the range of P.
cygnus with the size at first maturity ranging
from around 80 mm carapace length in the
northern areas of the fishery to around 95 mm
in southern areas (Morgan and Barker 1974).
No information is available on the size of first
maturity of males although it is probably similar
to that of females in the same area.

Mating takes place in July-August and the
female carries the black sperm packet (known
locally as a “tar-spot”) until November-Decem-
ber when the eggs are extruded and attach
themselves to the pleopods. Fecundity varies
linearly with the size of the rock lobster, a
100 mm carapace length female carrying 460 000
eggs (Morgan 1972). The eggs are carried on
the pleopods for up to 6 weeks, depending on
the water temperature, before hatching into the
first phyllosoma larvae that peak during Decem-
ber and January.

Larvae and juveniles

Studies of the rock lobster larvae have been
directed mainly towards understanding their
movements in space and time and their ecology.
Complementary investigations have also been
made to measure the numbers of larvae that
return to the coast at the end of the planktonic
phase of the life cycle.

The newly hatched larvae or phyllosoma, rise
up from the bottom, concentrate at the surface
and become part of the plankton. The body of
the phyllosoma is fiat and leaflike with long
legs and protuberent eyes. As there is no
pigment or calcium in the skeleton, the body is
colourless and transparent. The phyllosoma
larvae swim actively toward a weak light source
(Ritz 1972b) by beating their “feathery” legs
and most movements are directed to rising in
the water column since cessation of beating
results in a slow descent. The phyllosoma
larvae undergo a daily vertical migration in the
upper part of the water column, rising towards
the surface of the water toward dusk and then
descending to lower depths as dawn approaches
(Ritz 1972a). In their early stages they are

46

Journal of the Royal Society of Western Australia, Vol. 62, Parts 1-4, 1979.

transported offshore into the Indian Ocean by
the effects of wind-driven surface transport
acting on the larvae while they are at the
surface at night (Chittleborough and Thomas
1969).

The larvae remain in the phyllosoma stages
for 9 to 11 months during which time at least
9 phyllosoma stages are recognisable although
the number of actual moults is considerably
higher than this (George and Cawthorn 1962,
Chittleborough and Thomas 1969, Braine et al.
in press). The first phyllosoma stage is less
than 2 mm in length and because they are
almost transparent, are only just visible to the
human eye; by the last phyllosoma stage, they
are about 35 mm long.

The full extent of the larval distribution is
not known but significant numbers of larvae
have been caught during the intensive CSIRO
Fisheries larval research programme as far off-
shore as 1 500 km (Phillips et al. in press). Less
is known about the actual return paths of the
larvae to the coast of Western Australia.
Although many larvae are lost during the long
period in the plankton, the survivors are eventu-
ally transported back to the edge of the contin-
ental shelf of Western Australia by the ocean
circulation (Phillips 1977), the greater number
arriving between August and December each
year.

From this point the larval cycle is completed
by the transparent puerulus stage which is
similar in shape to the adult. The puerulus
stage metamorphoses from the last phyllosoma
stage just on or just beyond the continental
shelf and then actively swims across the shelf
and settles in the shallow reef areas along the
coast (Phillips et al. 1978). Collecting devices
made of artificial seaweed (Phillips 1972) have
been designed specially to catch the settling
puerulus and by their use, the times and relative
densities of settlement from year to year have
been able to be compared (Chittleborough and
Phillips 1975, 1977). Settlement follows a lunar
periodicity with catches being largely confined
to the new moon period (Phillips 1975).

The settling puerulus moults into a small pig-
mented rock lobster about 3 cm long, similar to
the adult. These juvenile rock lobsters remain
in the shallow coastal reefs (depths of 1-20 m)
for 3 to 6 years before migrating out into
deeper areas on the continental shelf (Chittle-
borough 1974d) .

Population biology

Much of the research effort which has been
expended on the western rock lobster has been
on those aspects of its population biology which
are of direct importance in making decisions
which affect the management of the fishery.
Apart from the routine collection of catch and
fishing-effort data for use in surplus-yield models
these data have included (a) estimates of growth
and mortality rates for use in various dynamic
pool models of the fishery, (b) information on
reproductive dynamics for assessing the likely
impact of fishing on the reproductive potential

of the stock and (c) various other biological
data which have been used to answer more
specific regional problems.

The “whites”

P. cygnus is unique among the world’s com-
mercially fished rock lobster stocks in that a
significant part of the commercial fishery is
based on pale coloured, newly moulted migratory
animals locally known as “whites”. Studies by
George (1958b) established that these “whites”
are all immature animals and move offshore
from the shallow water reefs (where they have
spent their early juvenile life) to deeper waters.
This late November migration may take the
animals on a journey of 25 km or more to an
area which presumably is better suited than the
inshore areas for the later release of the plank-
tonic larvae. Since the “whites” are newly
moulted animals their food requirements are
high (Chittleborough 1975) and consequently
their catchability by baited pots is likewise high
(Morgan 1974b). This results in high catch
rates being achieved by the commercial fisher-
men in late November and December and it is
during this 6 week period that about one third
of the annual catch is taken.

Growth

Since, like all other Crustacea, P. cygnus
grows by moulting, the growth process may be
considered to be a combination of two factors —
the moult increment (the size increase at each
moult) and the intermoult period (the time
between moults). Field-tagging studies invari-
ably are difficult to interpret in this light since
the observed growth between release and recap-
ture will consist of the unmeasured contributions
from these two growth processes. Studies in
aquaria however have been more fruitful and
have enabled meaningful interpretation of field-
growth studies.

The intermoult period increases with size in
P. cygnus and for animals held in aquaria at
25 °C the intermoult period is about 100 days
for an animal of 76 mm carapace length
(Chittleborough 1974d). Chittleborough (1976a)
and Morgan (1977a) have shown that moult
increment increases with age until the second
year of post-larval life, then remains constant
during the third year and then decreases with
increasing age. The combination of these two
processes results in a grown-in-length curve
which approximates the von Bertalanffy (1938)
curve and this curve has been used in the
biological models of the fishery (Morgan 1977a).

Although the “average” growth curve has been
used in biological models, growth rates in the
natural population exhibit large variations as a
result of the influences of limited food resources,
high densities of rock lobsters (Chittleborough
1976b) and temperature variations (Morgan
1977a). In addition, animals held in aquaria
were shown to exhibit a smaller moult increment
with a mild deficiency of oxygen while frequency
of moulting was reduced when animals were
held in isolation (Chittleborough 1975).

47

Journal of the Royal Society of Western Australia, Vol. 62, Parts 1-4, 1979.

Mortality

Mortality rates in P. cygnus may be considered
in two parts, those produced by fishing opera-
tions (the fishing mortality rate, F) and those
produced by natural causes such as predation
(the natural mortality rate, M). Since, like all
other crustaceans, P. cygnus cannot be individu-
ally aged, the usual methods of fisheries popu-
lation dynamics of comparing the abundance of
age classes to calculate F and M cannot be used
and so alternative methods have been necessary.
These have included direct estimation of M
from tagging studies (Chittleborough 1970,
Morgan 1974a) and the use of length/frequency
data to estimate F and M (Morgan 1977a).
These methods have indicated an annual value
of M of about 0.2 (equivalent to 18% mortality
each year) and a value of F of about 1.0
(equivalent to 63% mortality each year) for
recent years. Thus, mortality due to fishing
comprises some 80% of the total mortality
suffered by legal-size rock lobsters.

Management of the fishery

Since the start of the commercial fishery in
1944/45, both the catch and the amount of
fishing effort expended to take that catch have
increased dramatically (Table 1). This has
resulted in an inevitable decline in the catch
per unit of effort. During the 1940s and 1950s
management of the fishery was by means of a
legal minimum length only, since the amount
of fishing effort was relatively small and the
catch per unit of effort was high enough for
those engaged in the industry to achieve a
satisfactory return from their fishing activities.
In 1961, the Western Fisheries Research Com-
mittee met under the chairmanship of the then
Director of Fisheries to co-ordinate the research
efforts of State and Commonwealth programmes
to assist in management not only of rock lobsters
but of all the State’s fishery resources.

As fishing effort further increased and catch
per unit of effort further decreased it became
apparent from the analyses of the available
catch and effort data that more stringent
management measures would be necessary to
achieve the dual aims of providing a near maxi-
mum total catch and of providing a reasonable
return (in terms of catch per unit of effort)

to the individual operators in the fishery. The
most direct way in which this could be achieved
was by a limitation of effort policy and it was
this strategy which was adopted by the State
Government in 1962. The methods used to
limit fishing effort at that time were a restriction
on the number of vessels entitled to fish for
rock lobsters and a limit on the number of pots
each vessel was entitled to use. This pot alloca-
tion was based on boat length and was fixed at
3 pots per foot of boat length with a maximum
of 200 pots per boat.

Later, a boat replacement policy was intro-
duced which restricted a replacement vessel to
the same size as that vessel which was being
replaced. This measure prevented the number
of pots used being increased by the building of a
larger vessel. The effect of this legislation was
to reduce fishing effort from about 10 7 pot lifts
in 1962/63 to about 7 x 10 (i pot lifts in 1963/64.
Total catch decreased somewhat from 9.3 x 10 6 kg
in 1962/63 to 8.1 x 10 ,; kg in 1963/64 but sub-
sequently recovered to about 8.7 x 10 fi kg by
1966/67. Catch per pot lift improved slightly
from an average of 1.04 kg per pot lift for the
3 years prior to the introduction of the legis-
lation to 1.09 for the 3 subsequent years. How-
ever, the most important effect was that the
rapid escalation of the fishing effort of the late
1950s was halted.

Since the introduction of this limited entry
policy to control fishing effort, total catches
have remained stable at about 8 x 10 r> kg
although because of increased efficiency of the
individual operators in the industry, fishing
effort had by 1974 again increased to levels
similar to those immediately prior to 1962.
Further reduction in fishing effort was indicated
since subsequent analysis of fisheries data and
application of fisheries models confirmed the
concern experessed in 1962 regarding the effect
on catches of high levels of fishing effort. In
addition concern was expressed about the effect
of this high level of effort on the reproductive
potential of the stock.

The limited entry nature of the fishery, how-
ever, has posed some problems in the selection
of the most suitable methods to be used to
achieve further fishing effort reduction. One
of these problems is the increased capitalisation

Table 1

Average catch and fishing effort for the Western Australian rock lobster fishery , 1944-1975

Year

Average catch
(kg x 10 6 )

Average No. pot
lifts (x 10 6 )

Catch per
pot lift

Average No. of
men employed

1944/45-1948/49

0-994

0-618

1-608

109

1949/50-1953/54

3-547

2-284

1-553

402

1954/55-1958/59

5-805

4-172

1-391

734

1959/60-1963/64

8-541

7-815

1-093

1 378

1964/65-1968/69

8-434

7-647

1-103

1 350

1969/70-1973/74

7-338

9-407

0-780

1 668

48

Journal of the Royal Society of Western Australia, Vol. 62, Parts 1-4, 1979.

of the industry brought about by the fact that
the fishing licence (i.e. the right to fish) has
acquired a considerable monetary value. This
makes the reduction of boat or pot numbers an
almost prohibitively expensive exercise either for
Government or industry. Alternative strategies
which effect each individual equally are there-
fore necessary and some of the options which
have been examined are changing the legal
minimum size and implementing additional
closed seasons. In fact a 6 week additional
closure has been implemented on a trial basis
for the 1977/78 season. The problems of effort
reduction in a highly capitalised, limited entry
fishery such as the rock lobster industry have
not been fully resolved but solutions will need
to be found if increasing efficiency continues
to result in increasing fishing effort.

Although the Western Australian rock lobster
fishery stands as one of the few examples of a
well-managed fishery, those management tech-
niques that have been chosen have themselves
resulted in a new set of problems. Present
research is addressing itself to the resolution
of these new problems.

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Physiology and communication

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Technology

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741-748.

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4-8.

Chittleborough, R. G. (1974b). — Development of a tag
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Chittleborough, R. G. (1976b). — Breeding of Panulirus
longipes cygnus George under natural and
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Freshwat. Res. 27 : 499-516.

Griffiths, F. B. and Rimmer, D. W. (1977). — Description
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Oceanogr. Res. 94.

Hindley, J. P. R. and Penn, J. W. (1975). — Time lapse
photography of decapod crustaceans : A

comparison of white and infra-red illum-
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281-285.

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sorters. Fishery Industry News Service 6:
35-36.

Phillips, B. F. (1972). — A semi-quantitative collector of
the puerulus larvae of the western rock
lobster Panulirus longipes cygnus George
(Decapoda; Palinuridae). Crustaceana 22:
147-154.

Phillips, B. F. and Rimmer, D. W. (1975). — A surface
plankton sampler for the larval stages of
the western rock lobster. Aust. J. Mar.
Freshwat. Res. 26: 275-280.

Rimmer, D. W. and Phillips, B. F. (1978). — A pump net
sampler to examine the spatial distribution
of rock lobster larvae. CSIRO Aust. Div.
Fish. Oceanogr. Rep. 97: 1-18.

Sheard, K. (1950).— Care in the Handling of Crayfish.

West. Aust. Fish. Dept. Fisheries Bulletin 3.
Sheard, K. and Dicks, H. G. (1949). — Skin lesions among
fishermen at Houtman’s Abrolhos, Western
Australia, with an account of Erysipeloid of
Rosenbach. Rep. Med. J. Aus., 11: 352-354.

Wright, D., Yewers, P. and Softly, A. (1974). — Capture
and maintenance of some marine animals.
J. Inst. Anim. Tech. 25: 55-65.

Popular articles

Bowen, B. K. and Chittleborough, R. G. (1970). — Main-
taining the catch. Western Australian
Department of Fisheries and Fauna Pert.*'
Information Booklet.

George, R. W. (1966). — Marine Crayfish or Spiny Lobsters
of Australia. Australian Fisheries Newsletter,
May 1966.

George, R. W. (1978). — The tale of Rocky Lobster.
Your Museum 6: Jan. 1978.

Phillips, B. F. (1975b). — The World of the Rock Lohs+''-
Aust. Nat. Hist. 18: 305-307.

51

Journal of the Royal Society of Western Australia, Vol. 62, Parts 1-4, 1979, p. 53-64.

Aspects of Science in Western Australia 1829-1979.

A volume to commemorate the 150th Anniversary of the founding of Western Australia.

The breeding seasons of birds in south-western Australia

S. J. J. F. Davies

CSIRO Division of Wildlife Research, Helena Valley, W.A. 6056

Abstract

Data are presented on the dates when 2 703 clutches of eggs were laid by birds
breeding in south-western Australia, defined as that part of Australia west of the
Western Australian border and south of 26 °S latitude. In all except two species,
autumn breeding was rare and irregular. Over 90% of the clutches were laid
between July and December inclusive. Within that overall generalisation, some more
specific trends were evident.

Most species tended to lay earlier in the inland than in the south-western corner,
but the trend was reversed in two species. Species taking nectar as a food source
tended to breed before those taking grain. Ground insectivores bred earlier than
aerial insectivores. Raptors and water birds tended to breed later than other groups.

Consideration of data for individual species showed that although each species
had a characteristic breeding period, it varied regionally and from year to year in
the same place. No good correlation emerged between measurement of rainfall and
temperature and the onset or duration of breeding, although these factors appeared
to exercise some control in some species. Rather, it seems that the timing of the
birds’ breeding season relates directly to the abundance of the birds’ food supply
and may be controlled more by the food available for parents than by potential
food availability for the young.

It is pointed out that the concentration of breeding in south-western Australia
into the period from July to December implies that resources are most abundant then.
As a consequence, much more serious effects on the avifauna might be expected from
burning in the spring than in the autumn, because it would destroy the resources
which should provide food for the birds for the following twelve months.

Introduction

The time when birds breed in south-western
Australia, defined as that part of the continent
south of latitude 26 °S and west of the Western
Australian border, has attracted attention for
many years. Carnaby (1946) and Serventy
(1946) noted that although birds of south-west-
ern Australia normally bred in the spring,
autumn nesting was observed in some years.
(The terms spring and autumn refer to the sea-
sons as understood in the Southern Hemisphere;
spring is September to November, autumn is
March to May). They speculated that such out
of season nesting could be caused directly by out
of season rainfall (with which it had been ob-
served to correlate) or by an increase in food
supply consequent upon the rainfall.

Further observations were published by Carn-
aby (1954), Robinson (1955) and Sedgewick
(1955) which provided abundant evidence of a
spring breeding period in the far south-west and
occasional autumn breeding in northern areas.
Following Keast and Marshall’s (1954) publica-
tion of histological evidence of gonad response
to rainfall in Australian arid-zone birds, Ser-
venty and Marshall (1957) published detailed
observations of two autumn breeding episodes

in south-western Australia. Their data showed
that the gonads of many species enlarged after
cyclonic rains in March 1953 and February 1955,
that the males were stimulated more than the
females, that some species naturally laid eggs
and reared young in the autumns following the
cyclones, and that the incidence of this autumn
breeding was greater in the northern parts of
south-western Australia than in the more south-
westerly parts. Serventy and Marshall proposed
that the occurrence of spring breeding in south-
western Australia during a period of increasing
day length was probably coincidental. They
considered that the critical stimuli to reproduc-
tion are probably environmental conditions that
arise after rainfall and with the increasing tem-
peratures that accompany the lengthening
photo-period. They pointed out that low tem-
peratures appeared to inhibit breeding and that
the amount of unseasonal breeding might be
related to the degree of beneficial change to the
environment that followed the rain. They con-
sidered that these effects could not be observed
in all birds, notably not in sea birds, and con-
sidered the possibility that the breeding response
might be generated by a sudden improvement in
food supply consequent upon the rain, although

53

Journal of the Royal Society of Western Australia, Vol. 62, Parts 1-4, 1979.

they noted that the response was more rapid
than they would have expected an increase in
availability of food to be.

Little more was published about the breeding
seasons of birds in south-western Australia for
some years until Nix (1976) in an analysis of
the movements and breeding of birds in Aus-
tralia and New Guinea also considered the role
of food supply in the control of the timing of
breeding. He simulated on a computer the
growth periods of plants and showed that these
occurred at different times in different parts of
Australia. He suggested that the factors con-
trolling the timing of birds’ breeding seasons
would regulate them to coincide with the maxi-
mum food abundance. He suggested that the
breeding seasons of insectivores, nectarivores,
frugivores and graminivores would succeed each
other just as the stages of growth of the plants
succeeded each other. Taking data from Ford
and Stone, (1957) who published records of
breeding from Kellerberrin, he confirmed that
south-western Australian species have a major
breeding season in late winter and early spring
which is the period most favourable for plant
growth. His model showed that a smaller
episode of plant growth occurred in the autumn
and noted Serventy and Marshall’s (1957)
observations of autumn breeding.

The results of these earlier studies have dem-
onstrated that most breeding in birds in south-
western Australia occurs in the spring, but that
autumn breeding does occur sometimes, associ-
ated with unseasonal autumn rain. The authors
have universally accepted that an ultimate factor
controlling the timing of breeding must be the
birds’ food supply but the early speculation that
an important proximate factor timing the breed-
ing episode was rainfall itself has gradually lost
ground in favour of the speculation that the
environmental effects of the rainfall may be
more important. In addition, Nix (1976) has
nostulated a sequence of breeding that relates
to the sequence of plant growth.

Three recent studies of single species are now
available and show trends in individual species
more clearly. Nicholls (1964, 1974) has shown
that the wild individuals of the Silver Gull, Larus
novaehollandiae, can breed successfully in both
autumn and spring of the same year, and Saun-
ders (1977) has shown the same thing for wild
individuals of the Red-tailed Black Cockatoo,
Calyptorhynchus magnificus. Davies (1977) has
shown that although Zebra Finches, Poephila
guttata, breed mainly in the spring in the north-
ern part of south-western Australia as defined
here, they do breed in the autumn if summer
rains have brought up a crop of their main food,
Aristida contorta. In spring, successful breeding
correlates with high winter body weights and
therefore by implication with a good food supply
rather than with heavy rainfall.

Other data presented here comes from the Nest
Record Scheme of the Royal Australasian Orni-
thologists Union and from the records of myself
and my colleagues. It confirms the trends
already reported and expands the numbers of
species for which details can be given so that a

reasonable comparison can be made between the
succession of birds breeding and the sequence of
plant growth, as well as allowing further inter-
pretation of the role of rainfall and temperature
in timing the breeding episodes.

Methods and material

The collection of data in the Royal Australa-
sian Ornithologists Union Nest Record Scheme
has been examined and 1 298 cards found which
provide information about the dates when birds
breed in south-western Australia. Cards were
only accepted if they enabled the calendar month
in which eggs were in the nest to be determined.
If this spanned two months, the earlier was
chosen. In all cases, conservative estimates of
the egg-laying and incubation period were used,
for these are still unknown with respect to many
Australian birds. Personal records of 163 nests
with eggs at Mileura, Western Australia, have
been included with the RAOU nest record data.

Data on three species, the Zebra Finch, the
Yellow-rumped Thornbill, Acanthiza chrysor-
rhoa, and the Black-faced Woodswallow, Arta-
mus cinereus, were obtained from the original
notebooks of D. L. Serventy and relate to obser-
vations made at Grass Valley, York, from 1959
to 1963, referred to in Oksche et al. (1963).

Observations on Black-faced Woodswallows
nesting at Manmanning, on Splendid Wrens,
Malurus splendens, nesting at Gooseberry Hill
near Perth and on the Galah, Cacatua rosei-
capilla, have been contributed by Ian Rowley,
and data on the nesting of White-tailed Black
Cockatoos, Calyptorhynchus baudinii, in the
Western Australian wheatbelt, have been contri-
buted by Denis Saunders. In all cases, records
have only been accepted if the calendar month
in which there were eggs in the nest could be
unequivocally determined.

In the analysis of these records involving com-
parisons amongst them, statistical methods are
difficult to use, both because nest searching can-
not be considered to be non-random and because
the coverage of the area under consideration is
not even. For example, there are abundant
records from a few centres where keen orni-
thologists reside such as Woodanilling, Mileura,
the Nullarbor Plain and Manjimup. This problem
affects all studies of when birds breed, e.g. Frith
and Davies (1961) and Lavery et al. (1968), and
it is only when the raw data show overwhelm-
ing trends that reasonable certainty can be
assumed. In the present study, D. L. Serventy ’s
data from York are of great value, for the area
was visited regularly each three weeks and
searched systematically for nests on each visit.
The trends showing up in his results should,
therefore, be free from bias due to non-random
searching and provide an accurate picture of the
timing of breeding of these three species in the
York area.

In order to analyse regional trends within
south-western Australia itself, regional bound-
aries used by the Commonwealth Bureau of
Meteorology, in their published statistics, have

54

Journal of the Royal Society of Western Australia, Vol. 62, Parts 1-4, 1979.

been adapted and are shown in Figure 1. The
Bureau of Meteorology’s south-west divisions 9
and 9A correspond to the south-west division
of Figure 1; its south-west divisions 8, 10 and
10A correspond to the wheatbelt division of
Figure 1 and its divisions 6 (part), 7 (part), 7A,
11, 12 and 13 (part) correspond to the inland
division of Figure 1. These boundaries do not
correspond exactly with mean annual isohyets
as shown, for example, by Nix (1976) but are
quite close to the 600 mm and 350 mm isohyets,
and conveniently divide the forested south-west
from the cleared wheatbelt and the arid interior.

Figure 1. — Map of south-western Australia showing the
three regional sub-divisions used in this paper.

Results

RAOU Nest Record Scheme and personal records
from Mileura

Table 1 shows an analysis of the breeding sea-
son data available for species represented by less
than 10 records. The species have been grouped
by their broad ecological niche, as has become
conventional in regional studies of breeding sea-
sons (Frith and Davies 1961, Lavery et al. 1968).
The species included under each heading are
listed in Appendix I. Table 1 indicates a pre-
dominantly spring breeding season: 31% of

records recorded eggs in the nest in September,
68% recorded eggs in the nest in the three spring
months, and 94% recorded eggs in the nest be-
tween July and December. The sequence of nest-
ing insectivores, nectarivores and graminivores
follows that hypothesised by Nix (1976), except
that aerial insectivores are considerably later
than ground insectivores. Table 2 gives data for
species represented by 10 to 19 records. Again
there is evidence of a predominantly spring
breeding season: 32% of records recorded eggs
in the nest in September, 65% recorded eggs in
the nest in the three spring months, and 93%
of the records were of eggs in the nest between
July and December. Of the individual species,
those that had eggs in the nest between January
and June were the aquatic Little Grebe, Tachy-
baptus novaehollandiae , and migrant Clamorous
Reed Warbler, Acrocephalus stentoreus, both of
which nested late, and a number of species in
which odd individuals nested early, the White-
faced Heron, Ardea novaehollandiae (2), Wood
Duck, Chenonetta jubata (1), Whistling Kite,
Haliastur sphenurus (1), Wedge-tailed Eagle,
Aquila audax (4), Crested Pigeon, Ocyphaps
lophotes (3), Richard’s Pipit, Anthus novaesee-
landiae (1), Black-faced Cuckoo-shrike, Coracina
novaehollandiae (2), White-fronted Chat, Eph-
thianura albifrons (2), Australian Magpie lark,
Grallina cyanoleuca (1), and Grey Butcherbird,
Cracticus torquatus (1). In some cases these
nests were earlier than the bulk of the popula-
tion and others may best be classified as autumn
nesting attempts. Tables 1 and 2 indicate a
clear tendency for species of birds in south-
western Australia to nest from July to Decem-
ber. This trend is evident even though regional
differences and differences between individual
years cannot be taken into account in these data.

Table 1

Breeding seasons of birds in south-western Australia ( species with less than 10 records')

J

F

M

A

M

J

J

A

S

O

N

D

Totals

Aerial Insectivores ....

1

10

4

4

3

22

Ground Insectivores ...

1

2

i

15

28

46

29

20

4

146

Nectarivores ....

2

i

4

16

19

13

9

1

65

Graminivores

4

2

2

1

16

17

8

6

1

57

Omnivores

1

2

9

4

16

Shore and sea birds ....

1

2

4

9

4

11

4

35

Raptors

1

14

12

3

30

Water birds

3

5

5

2

10

9

34

Totals

6

4

2

1

5

4

22

80

124

72

63

22

405

55

Journal of the Royal Society of Western Australia, Vol. 62, Parts 1-4, 1979.

Table 2

Breeding seasons of birds in south-western Australia ( species with 10-19 records)

J

F

M

A

M

J

J

A

S

O

N

D

1

Main data from

Little Grebe

Tachybaptus novaeholland-
iae

White-faced Heron

4

2

3

1

4

Manjimup (8), Wooroloo (4)

1

1

3

4

2

Woodanilling (5)

Ardea novaehollandiae
Mountain Duck

8

6

1

Woodanilling (1 1)

Tadorna tadornoides
Black Duck

2

7

1

Yunderup (1), Benger (4)

Anas super ciliosa
Grey Teal

2

8

2

3

Woodanilling (6)

Anas gibberifrons
Wood Duck

3

2

7

1

!

Woodanilling (6)

Chenonetta jubata
Whistling Kite

1

7

2

Woodanilling (7)

H alias tur sphenurus
Wedge-tailed Eagle

4

3

4

3

,

Mileura (9)

Aquila audax
Brown Falcon

2

8

2

Nullarbor Plain (8)

Falco berigora

Little Button-quail

8

2

1

1

1

Rawlinna (12)

Tur nix velox
Crested Pigeon

1

2

5

3

Mileura (4)

Ocyphaps lophotes
Galah

12

9

Kathleen Valley (9)

Cacatua roseicapilla
Port Lincoln Parrot

3

5

2

Cunderdin (3), Perth (2)

Barnardius zonarius

Tawny Frogmouth

4

6

1

Mileura (2). Nullarbor Plain

Podargus strigoides

(2), Woodanilling (2)

Sacred Kingfisher ....

10

Katanning (3), Woodan-

Halcyon sancta
Richard’s Pipit

filing (2)

1

3

4

5

2

2

Woodanilling (5), Nullarbor

Anthus novae seelandiae

Plain (5)

Black-faced Cuckoo-shrike

1

1

3

4

5

2

1

Woodanilling (4), Nullarbor

Coracina novaehollandiae

Plain (3)

Ground Cuckoo-shrike

5

7

Nullarbor Plain (8)

Coracina maxima
White-winged Triller

4

4

4

1

Woodanilling (7)

Lalage sueurii
Grey Shrike-thrush

1

2

6

3

Nullarbor Plain (5), Wood-

Colluricincla harmonica

anilling (3)

Clamorous Reed Warbler

8

3

7

Manjimup (18)

Acrocephalus stent or eus
Brown Songlark

9

1

Nullarbor Plain (10)

Cinclorhampus cruralis
Varied Sittella

3

2

3

2

Woodanilling (7)

Daphoensitta chrysoptera
Yellow-throated Miner

2

5

12

2

Woodanilling (7), Cunder-

Manorina flavigula

din (5)

Crimson Chat

4

2

7

5

Nullarbor Plain (14)

Ephthianura tricolor
White-fronted Chat

1

1

7

2

Woodanilling (7)

Ephthianura albifrons
Magpie Lark

1

1

5

4

2

Woodanilling (5), Miling (3)

Grallina cyanoleuca
Masked Woodswallow

4

2

1

10

Nullarbor Plain (10)

Artamus personatus
Dusky Woodswallow

8

9

1

Woodanilling (9), Manjimup

Artamus cvanopterus

(5)

Grey Butcherbird

1

1

17

1

Coolup (10)

Cracticus torquatus

Totals

12

4

4

2

2

7

41

92

134

48

56

17

Grand total: 419

Table 3

(a) — Grouped breeding seasons — all birds with more than 20 records; summary by months

District

J F

M

A

M

J

J

A

S

O

N

D

Totals

s.w

1

1

7

4

25

33

38

21

11

141

Wheatbelt

1

2

10

6

18

31

32

30

14

6

150

Inland

2

15

12

20

14

31

87

127

19

37

364

Totals

1 3

15

14

31

27

53

143

192

87

72

17

655

56

Journal of the Royal Society of Western Australia, Vol. 62, Parts 1-4, 1979.

(b ) — Breeding seasons grouped by districts; species with more than 20 cards

Species

District

J

F

M

A

M

J

J

A

S

O

N

D

Totals

Willie Wagtail

SW

1

1

8

3

3

16

Wheatbelt

5

10

2

2

19

Inland

i

1

11

2

15

Red-capped Robin

Wheatbelt

5

6

5

5

21

Inland

6

1

7

Black-faced Woodswallow

Wheatbelt

2

6

1

9

Inland

13

2

6

21

Welcome Swallow

SW

8

13

9

6

36

Inland

i

3

2

1

7

Silvereye

SW

1

5

3

2

2

13

Wheatbelt

2

6

8

Magpie

SW

9

10

4

23

Wheatbelt

9

7

3

19

Black Swan

Wheatbelt

2

3

2

7

Inland

4

3

ii

18

Banded Plover

Wheatbelt

l

4

6

2

2

1

16

Inland

1

i

6

8

16

Australian Dotterel

Wheatbelt

1

l

6

2

1

1

12

Inland

1

2

4

8

2

1

18

Coot

SW

5

1

4

9

9

28

Red Wattlebird ....

SW

1

1

1

2

6

3

14

Raven

Wheatbelt

4

6

6

3

19*

Kestrel

Inland

5

50

4

3

62

White-browed Babbler ....

Inland

1

3

2

1

3

3

7

3

23

Zebra Finch

Inland

1

10

5

2

6

7

16

13

12

22

94

Little Crow

Inland

3

6

41

26

76

Totals

3

15

14

29

22

47

131

186

77

70

17

6

617

*One card from the SW.

For those species represented in the records
by 20 or more cards, regional differences can be
examined. The data are presented in Table 3.
Table 3(a) presents the grouped results: 29%
recorded eggs in the nest in September, 64%
recorded eggs in the nest in the three spring
months, and 86% recorded eggs in the nest be-
tween July and December. Looking at regional
differences, the percentage of records of eggs in
the nest in September was 23, 21 and 35 for the
south-west, wheatbelt and inland respectively.
This was the highest percentage of any month
for the inland but in the wheatbelt and south-
west, the spread was more even over August,
September and October and in the south-west a
slightly higher percentage occurred in October
(27%) although the difference is small.

The data for individual species with sufficient
records for at least one region to be considered
separately are shown in Table 3(b), and the
trends are easier to see. The Willie Wagtail,
Rhipidura leucopbyrs, Red-capped Robin, Pet-
roica goodenovii, and Black-faced Woodswallow
tend to nest earlier in the inland than in either
the wheatbelt or the south-west. The Australian
Magpie, Gymnorhina tibicen and Silvereye, Zos-
terops lateralis, do not show differences between
wheatbelt and south-west although the Silvereye
appears to have a more extended breeding season

in the south-west than the wheatbelt. On the
other hand, both the Banded Plover, Vanellus
tricolor and the Australian Dotterel, Peltohyas
australis, tend to nest a little earlier in the
wheatbelt than the inland. There are insuffici-
ent data from the other species to compare
regions.

A second important phenomenon begins to
emerge in Table 3. Some species characteristi-
cally breed both between January and June and
between July and December whilst others breed
only between July and December. Thus all the
nests of the Willie Wagtail, Red -capped Robin,
Black-faced Woodswallow, Australian Magpie
and Nankeen Kestrel, Falco cenchroides, occur
between July and December. A few Australian
Ravens, Corvus coronoides, and Little Crows,
Corvus bennetti, nested in June and one Silvereye
and a few Coots, Fulica atra, were recorded with
eggs in the nest in January, but otherwise these
four species only nested between July and Dec-
ember. In other species some nests were recorded
between January and June but in these cases
there were fewer nests between January and
June than between July and December. These
species were (figures in parentheses compare
nests in first and second half of the year) :
Banded Plover (7:25), Welcome Swallow, Hirundo
neoxena (1:42), Red Wattlebird, Anthochaera

57

Journal of the Royal Society of Western Australia, Vol. 62, Parts 1-4, 1979.

Table 4

Breeding seasons ( number of nests with eggs ) of three passerines at York over five seasons {data from D. L. Serventy)

Species/Year

No. of
Visits

M

J

J

A

s

O

N

D

J

F

M

A

M

Totals

Autumn

nests

Zebra Finch Poephila gut-

tata —

1959-60

14

1

11

4

1

1

4

1

23

7

1960-61

17

1

3

2

2

7

3

1

4

23

15

1961-62

17

1

6

5

2

1

15

3

1962-63

17

1

3

2

1

1

4

3

15

9

1963-64

18

i

2

3

3

0

1

2

12

6

Totals

1

3

17

18

9

10

9

6

12

3

88

40

Yellow-rumped Thornbill

Acanthiza chrysorrhoa—

1959-60

14

2

4

1

7

1960-61

17

1

4

3

1

2

4

15

1961-62

17

2

4

3

3

12

1962-63

17

5

1

3

9

1963-64

18

1

1

2

Total

1

1

5

10

8

12

8

45

Black-faced Woodswallow

Artamus cinereus —

1959-60

14

1

1

1

1

4

1

1960-61

17

3

1

4

1961-62

17

1

4

4

9

1962-63

17

1

1

1963-64

18

0

Totals

....

-•

2

8

7

1

18

1

carunculata (2:12), White-browed Babbler,
Pomatostomus super ciliosus (7:16) and Zebra
Finch (24:70). Two species stand apart. The Aus-
tralian Dotterel appears to be predominantly an
autumn and winter nesting species, both in the
wheatbelt and the inland and the Black Swan,
Cygnus atratus, has been recorded nesting in
the autumn in the inland and the late winter,
early spring in the wheatbelt. Further data col-
lected by Tingay et al. (1977) indicate that it
nests in the late winter in the south-west.

Other data

Grass Valley, York. — D. L. Serventy’s data are
presented in Table 4 and show that each species
has a characteristic breeding pattern. The Zebra
Finch shows a large peak of nests with eggs in
October and November, and a smaller peak in
April. This resembles the data for that species
shown in Table 3, but the start of intense spring
nesting is earlier in the inland than in the
wheatbelt at York. On the other hand, summer
nests occur in the wheatbelt whereas they are
rare in the inland. Most nests of the Yellow-
rumped Thornbill were recorded in August, Sep-
tember and October and most of the Black-faced
Woodswallow nests were in November and Dec-
ember, in agreement with the trend shown in
Table 3(b) for this species to nest later in the
wheatbelt than at inland sites.

Differences between years are evident. In
1959-60 and 1963-64, the Zebra Finch had well
separated spring and autumn breeding seasons.
In 1961-62 breeding was over in February, and
the species effectively bred only in the spring.
On the other hand, in 1962-63, most of the
breeding was in the summer and autumn, and
in 1960-61 breeding was continual from Septem-
ber to April.

The total monthly rainfall at York is shown in
Figure 2. It is not easy to find direct correla-
tions between rainfall and the breeding of Zebra
Finches. The total nests found in the study
area each season correlates positively (Spear-
man’s r = +0.95; P< 0.05) with the total rain-
fall in January of the year in question, but it
is not easy to accept this as a valid correlation,
for in some years, e.g. 1961-62, the bulk of the
nesting is over by January. There seems to be
no clear association between the start or end
of the winter rain period and the start of the
breeding season, nor between summer rainfall
and the presence of summer breeding. In Figure
2, the mean monthly temperatures recorded at
Perth are shown. Although this locality is 100
km west of York, the trend shown in a compari-
son across years would be similar in the two sites.
There is no clear correlation between tempera-
ture and the pattern of breeding shown by the
Zebra Finches. It is likely, therefore, that the
birds are not responding directly to these en-
vironmental factors, although the timing of
breeding may be controlled by the indirect effects
of environmental changes.

The same statement may be made about the
breeding of the Yellow-rumped Thornbill and
the Black-faced Woodswallow. In both there
are differences between years but in neither do
these differences relate directly to differences
between rainfall and temperature patterns in the
corresponding years. Yellow-rumped Thornhills
bred early in 1960-61 and 1963-64 and in 1963-64
failed to breed at all in their normal spring
period. In comparison with Ford’s data from
Bibra Lake, Perth (Ford, 1963) for some of the
same years (1959 and 1961) this species bred one
to two months earlier at York than on the coast.
The Black-faced Woodswallows bred consistently

58

Journal of the Royal Society of Western Australia, Vol. 62, Parts 1-4, 1979.

1959 1960 1961 1962 1963 1964

1973 1974 1975 1976 1977 1978

Figure 2. — Environmental data for three sites at which breeding data were collected, (a) Monthly rainfall totals
for York, (b) Mean monthly temperatures for Perth. (c) Monthly rainfall totals for Cadoux. (d) Monthly

rainfall totals for Kalamunda.

in late spring, early summer, but in 1962-63 and
in 1963-64 only one nest was found in the study
area, and breeding did not seem to have taken
place.

Table 5

Breeding data from Rowley for two species of birds in south-western
Australia. ( Number of nests with eggs.)

Serventy’s data, therefore, indicate that in one
place, each species has a characteristic breeding
time, but that the precise timing varies from
year to year. Further, the changes in timing
with different years do not correlate between
species. Thus the Zebra Finch bred early in
1959-60 but late in 1960-61, whereas the reverse
was true for the Yellow-rumped Thornbill.

Manmanning. — Rowley’s data for the Black-
faced Woodswallow are presented in Table 5 and
show that breeding is a month earlier than for
the same species at York. Again there are dif-
ferences between individual years both in timing
and extent of the breeding period. The mean
monthly rainfall at Cadoux, 8 km north of Man-
manning is shown in Figure 2. There is no clear
correlation between differences in the timing and
extent of the breeding season in individual years
and differences in the pattern of precipitation in
corresponding years.

Month

Years Total

S O N D J

(a) Black-faced Woodswallow, Manmanning.

1973-74

3

7

3

2

15

1974-75

8

14

3

25

1975-76

3

12

15

1976-77

8

4

12

Totals

22

37

6

2

67

(b) Splendid Wren, Gooseberry Hill, Kalamunda.

1973-74

2

3

2

7

1974-75

T

8

3

3

1

16

1975-76

5

5

2

12

1976-77

7

3

1

1 1

1977-78

1

3

3

1

1

9

Totals ....

11

22

14

6

2

55

59

Journal of the Royal Society of Western Australia, Vol. 62, Parts 1-4, 1979.

Gooseberry Hill. — Rowley’s data for the Splen-
did Wren show large differences between years
in both the timing and extent of breeding. Rain-
fall figures for nearby Kalamunda are presented
in Figure 2 and again no correlation is evident.
There is a general trend for the species to nest
at the end of the wet winter, but detailed cor-
relations do not occur.

Galah. — Rowley’s data for the Galah are pre-
sented in Table 6. The data were collected at
three sites, Manmanning and Coomalloo in the
south-west division and Mileura in the inland

division. Galahs show considerable constancy
in the timing of the nesting season from year to
year and region to region. Where differences
between years occur, Manmanning and Coomal-
loo differ in the same direction. At both sites the
month in which most nests were started was
August in 1971 and 1974 and September in 1972
and 1973. The data from Mileura are not exten-
sive but do indicate a concentration of nesting
in the August-September period. Overall 90%
of the 581 Galah nests were started in August
or September.

Table 6

Breeding data from Rowley for the Galah from three sites in south-western Australia.

The figures indicate the number of nests started each month

Year

J FMAMJ JASOND

Total

Coomalloo Creek

1971

23 4 5

32

1972

7 15 4

26

1973

4 12 2

18

1974

2 23 4 1

30

Manmanning ....

1971

1 68 26 8

103

1972

16 67 11

94

1973

43 53 14

110

1974

88 28 6

122

Mileura ....

1971

1 14 6 2

23

1972

14 2

16

1974

5 2

7

Totals

4 305 219 53

581

White-tailed Black Cockatoo . — Table 7 presents
Saunders data for the breeding seasons of this
species at four different localities in south-west-
ern Australia. All, except Coomalloo Creek
(south-west) are in the wheatbelt division of the
State. Overall differences are apparent in the
timing of breeding. The most northerly locality,
Coomalloo Creek, is earliest even though it is in

the south-west division, and the most southern
locality, Moornaming, is latest, with Manman-
ning and Tarwonga in an intermediate position.
There are differences too between individual
years at each locality. Saunders (1979) dis-
cusses these differences and can find no clear
correlation between environmental variables and
the timing of breeding in this species.

Table 7

Breeding data from Saunders ( number of nests with eggs) of the White-tailed Cockatoo in four localities in south-western Australia

Year

M

J J

A

S

O

N

D

J

F

M

A

Total

Coomalloo Creek

1969

3

5

1

9

1970

10

13

5

28

1971

15

30

9

1

55

1972

25

37

3

65

1973

4

43

19

8

74

1974

13

37

13

1 1

1

75

1975

10

39

23

9

1

82

1976

2

27

28

7

64

1977

8

31

2

41

Manmanning

1969

1

2

2

1

6

1970

3

12

3

18

1971

12

8

2

22

1972

1 1

2

13

1973

3

5

6

14

1974

7

6

13

1975

4

6

10

1976

3

3

6

1977

0

Tarwonga

1969

1

4

1

3

9

1970

1

5

7

13

Moornaming

1969

2

4

6

1970

13

2

1

16

Totals

49

265

250

72

3

639

60

Journal of the Royal Society of Western Australia, Vol. 62, Parts 1-4, 1979.

Mileura . — Davies (1977) has presented breed-
ing data for the Zebra Pinch at this inland site
which may be compared with Serventy’s data
from York, a wheatbelt site. The pattern of
breeding exhibited by the species is similar, with
breeding taking place in the spring and to a
lesser extent throughout summer and autumn,
but varying in its precise timing from year to
year. There was some evidence that chicks
raised from autumn nestings survived less well
than those raised during spring nestings.

Discussion

The results presented above show that the
birds of south-western Australia tend to breed
between July and December, predominantly in
the spring months. Within that overall generalis-
ation, some specific trends are evident. Although
many species breed only in the spring, some also
exhibit autumn breeding activities, as discussed
at length by Serventy and Marshall (1957). In
most cases these autumn breeding episodes are
quantitatively smaller than the equivalent spring
ones, but for the Australian Dotterel throughout
the region, and the Black Swan in the inland,
autumn breeding appears to be the dominant
pattern. Regional trends also appear. In gene-
ral, species breed earlier inland than in the
wheatbelt or south-west. The two plovers are
interesting, however, in that both breed earlier
in the wheatbelt than the inland and the north-
ern populations of White-tailed Black Cockatoos
breed earlier than the southern ones.

Clear variations in the timing of breeding for
individual species from year to year, suggest that
photoperiod itself is not always a major deter-
minant. Serventy and Marshall (1957) have
already stressed this point. They consider that
the critical stimuli to reproduction are probably
environmental conditions that arise after rain-
fall and with relatively high temperatures that
accompany the lengthening photoperiod. They
also considered low temperatures to be important
as a breeding inhibitor.

South-western Australia experiences a winter
rainfall climate in which heavy rain is rare be-
tween November and March. The trend of
south-western species to breed between July and
December is, therefore, consistent with Serventy
and Marshall’s hypothesis, for in most years the
spring is the only period following heavy rain
that is also warm. Such a general hypothesis
cannot be expected to apply to all species and the
White-browed Babbler, Banded Plover and Aus-
tralian Dotterel all nest in the winter, a cold
wet period. Probably these ground-feeding in-
sectivorous birds find food with difficulty in dry
weather, but with much greater ease as the in-
sects become active after rain. They may, there-
fore. be responding either to the rain, or the
environmental effects of it or simply to the in-
creased food supply.

The year to year variation in the detailed data
given for the Zebra Finch, Yellow-rumped
Thornbill, Black-faced Woodswallow, Splendid
Wren, Galah and White-tailed Black Cockatoo
show that each species must be considered separ-

ately in relation to environmental factors and
that no simple relationships are evident between
rainfall and breeding response.

Marshall (1961) reviews work that suggests
that the timing of breeding episodes is correlated
with ensuring a good food supply for the young.
Much of the work up to that time sought to
establish correlations between the onset of breed-
ing and environmental variables that might pre-
cede such abundant food supplies for the young,
although they may not themselves be causally
related to increasing this food supply. More
recently, Perrins (1970) discussed the role of food
in the timing of breeding and concluded that the
evidence, although not yet overwhelming, was
tending to implicate the food supply of adults,
particularly of the females, as an important
proximate factor controlling the onset of birds’
breeding. Recent work by Jones and Ward
(1976) on the Red-billed Quelea, Quelea quelea,
suggests that its breeding episodes may be timed
more precisely by direct relationships between
the present food supply for potential parents
rather than the future food supply for young.

Some of the detailed studies of breeding in
south-western Australian birds that have been
published, show correlations between good
supplies of food for the parents and breeding
episodes. Nicholls’ (1964, 1974) studies of the tim-
ing of breeding of the Silver Gull showed that
there are eggs continuously on Carnac Island, off
Perth, throughout the autumn, winter and
spring, with little egg-laying in the winter. Very
little laying occurs in summer. The food sup-
plies of Silver Gulls in metropolitan Perth come
largely from the suburban rubbish tips and offal
sites adjacent to slaughter yards. These are
available throughout the year. Studies by
Nicholls of captive gulls showed that young
chicks hatched in the summer died of heat, and
if this effect operated in the wild, as it almost
certainly does, there would be strong selection
against gulls breeding in the summer, because
energy would be wasted in a regularly unsuccess-
ful breeding attempt. Silver Gulls, therefore,
breed throughout that part of the year when
chicks will survive. They have a nearly constant,
plentiful food supply and the lack of winter
egg-laying is probably a consequence of the
nearly simultaneous start of breeding by gulls
in April-May, that involves them in parental
duties for the next three months. The only
winter eggs would come from second breeding
attempts by failed breeders.

Saunders (1977) studying the Red-tailed Black
Cockatoo, showed that it bred in both spring and
autumn. This species has taken to eating the
fruits of an introduced weed, Emex australis,
almost exclusively and the double breeding sea-
son may also reflect an abundant and constant
food supply.

At Mileura, Davies (1977) was able to show
that Zebra Finches bred in warm periods, especi-
ally the spring, in years of effective summer
rainfall when the birds’ food supply of grass had
germinated, established and seeded well. In three
years studied there was little successful breeding
in one year and much successful breeding in two.

61

Journal of the Royal Society of Western Australia, Vol. 62, Parts 1-4, 1979.

In the year of little breeding the birds’ winter
weights were lower than in the other two years,
suggesting that the condition of the parents may
be a factor determining whether or not breeding
occurs. The Zebra Finch definitely did not
respond by breeding after any rainy episode. The
young hatched in the autumn appeared not to
survive as well as those hatched in the spring,
and from an evolutionary point of view may seem
unimportant. On the other hand, Zebra Finch
populations sustain great fluctuations, and poten-
tially have a very high rate of increase, so that
the survival of even a few birds might, in certain
years, make the difference between the survival
and recovery of the local population, or its total
extinction. In such circumstances even a small
autumn breeding episode could be important in
giving a depleted population additional recruits
for a spring-breeding period.

The Willie Wagtail (Table 3(b)) and Black-
faced Woodswallow (Tables 3(b) and 4) are both
aerial insectivores and tend to breed later than
the ground insectivores, the Yellow-rumped
Thornbill (Table 4) and the Red-capped Robin
(Table 3(b)). This appears to correlate well
with the activity of their prey, for whereas
ground-dwelling insects become active as the
soil wets with the first rains, the flying insects
are not abundant until the weather warms up
and the growth and resting stages metamorphose
and begin flying. The aerial insectivores are
correspondingly later in breeding than the
ground insectivores. Detailed studies of indi-
vidual species are needed to resolve this issue.

An interesting case is the White-tailed Black
Cockatoo (Table 5) which lays early in the spring.
It feeds on the fruits of Proteaceae which are
available at that time. However, a second factor
probably operates here, for the bird has a very
long incubation and nestling period (Saunders
pers. comm. 1978). Young are sometimes desic-
cated by the heat of summer if they have not
left the nest by the end of November, and there
would be in these cockatoos, as in the seagulls,
strong selective pressures against starting nesting
attempts late in the year. It is important to
view this restraint in perspective for the distri-
bution of these cockatoos is probably limited to
areas where there is sufficient food available
early in the year for them to start nesting in
time to get the young out of the nest before the
hot weather begins.

A clear assessment of the importance of the role
of parental food supply as opposed to environ-
mental variables in timing the initiation of
breeding episodes of south-western Australian
birds must await more data and preferably
experimental work for which there is now ample
field information to ensure unequivocal experi-
mental design. Nonetheless one important con-
clusion can be drawn from this survey. The
breeding seasons of south-western Australian
birds are concentrated between July and Dec-
ember and occur particularly in the spring. Be-
cause there seems to be a clear correlation be-
tween the time birds breed and an abundant
food supply, either for parents or young, it fol-
lows that the food resources on which these birds

depend are particularly abundant then, and this
abundance also provides a store on which they
will draw for the rest of the year. If this store
is destroyed, as it would be by burning off in the
spring, not only will their breeding effort be
jeopardised, but their store of future food will
also be damaged. Although Christensen and Kim-
ber (1975) have found that bird populations are
not greatly disturbed by burning, it is clear from
the present review that spring burning must be
much more damaging than autumn burning.
Further, as many birds feed on invertebrates, at
least part of this food store must represent a
peak of abundance of invertebate life, so that
burning in the spring would damage the produc-
tivity and perhaps the variety of the invertebrate
fauna even more severely than it would bird
populations.

Acknowledgements. — I am grateful to the RAOU Nest
Record Scheme for permission to consult its card
collection and to use the da' a abstracted. I am also
grateful to Ian Rowley, D. A. Saunders, D. L. Serventy,
C. A. Nicholls and P. de Rebeira for permission to use
data on laying dates collected by them. D. L. Serventy,
Ian Rowley and D. A. Saunders have kindly read and
criticised this paper in draft form. I am grateful to
P. de Rebeira for preparing the figures.

References

Carnaby, I. C. (1946). — Notes. W.A. Bird Notes 4: 11.

Carnaby, I. C. (1954). — Nesting seasons of Western Aus-
tralia birds. W.A. Nat. 4: 149-156.

Christensen, P. and Kimber, P. (1975). — The effects ol
prescribed burning on the flora and fauna
of south-west Australian forests. Proc. Ecol.
Soc. Aust. 9: 85-106.

Davies, S. J. J. F. (1977). — The timing of breeding of
the Zebra Finch Taeniopygia castanotis at
Mileura, Western Australia. Ibis 119: 363-372.

Ford, J. R. and Stone, P.S. (1957). — Birds of the Keller-
berrin/Kwolyin, district, Western Australia.
Emu 57: 9-21.

Ford, J. R. (1963). — Breeding behaviour of the Yellow-
tailed Thornbill in south-western Australia.
Emu 63: 185-200.

Keast, J. A. and Marshall, A. J. (1954). — The influence
of drought and rainfall on reproduction in
Australian desert birds. Proc. Zool. Soc.
Lond. 124: 493-499.

Jones, P. J. and Ward, P. (1976). — The level of reserve
protein as the proximate factor controlling
the timing of breeding and clutch-size in
the Red-billed Quelea, Quelea quelea. Ibis
118: 547-574.

Lavery, H. J., Sefton, D. and Bravery, J. A. (1963). —
Breeding seasons of birds in north-eastern
Australia. Emu 68: 133.

Marshall, A. J. (1961). — Food availability as a timing
factor in the sexual cycle of birds. Emu 50:
267-282.

Nicholls, C. A. (1964). — Double-broodedness in the Sil-
ver Gull, Larus novaehollandiae . W.A. Nat.
9: 73-77.

Nicholls, C. A. (1974). — Double-brooding in a Western
Australian population of the Silver Gull,
Larus novaehollandiae. Aust. J. Zool. 22 :
63-70.

Nix, H. A. (1976). — Environmental control of breeding,
post-breeding dispersal and migration of
birds in the Australian region. Proc. XVI
Int. Ornith. Cong.: 272.

Oksche, A., Farner, D. S., Serventy, D. L., Wolff, F.

and Nicholls, C. A. (1963). — The hypothal-
amo-hypophysial neurosecretory system of
the Zebra Finch, Taeniopygia castanotis.
Ziet. f. Zellforsch. 58: 845-914.

Perrins, C. M. (1970). — The timing of birds breeding-
seasons. Ibis 122: 243-255.

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Robinson, A. H. (1955). — Nesting seasons of Western
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W.A. Nat. 4: 187-192.

Saunders, D. A. (1977). — Red-tailed Black Cockatoo
breeding twice a year in the south-west of
Western Australia. Emu 77: 107-110.

Saunders, D. A. (1979). — The biology of the short-
billed form of the White-tailed Black
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prep.).

Sedgewick, E. H. (1955). — Nesting records — 1946 and
1947. W.A. Nat. 5: 46-47.

Serventy, D. L. (1946). — The pituitary gland and the
breeding cycle. W.A. Bird Notes 4: 10-11.

Serventy, D. L. and Marshall, A. J. (1957). — Breeding
periodicity in Western Australian birds
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185-187.

Grey-crowned Babbler
Little Grassbird
Splendid Wren
Red-winged Wren
White-winged Wren
Rufous-crowned Emu-
wren

White-browed Scrubwren

Redthroat

Weebill

Western Warbler
Broad-tailed Thornhill
Chestnut-rumped
Thornhill

Samphire Thornbill
Southern Whiteface
White-browed Treecreeper
Rufous Treecreeper
Striated Pardalote

Pomatostomus temporalis
Megalurus gramineus
Malurus splendens
Malurus elegans
Malurus leucopterus

Stipiturus ruficeps
Sericornis frontalis
Sericornis brunneus
Smicrornis brevirostris
Gerygone fusca
Acanthiza apicalis
Acanthiza uropygialis

Acanthiza iredalei
Aphelocephala leucopsis
Climacteris affinis
Climacteris rufa
Pardalotus striatus

Omnivores

Appendix I

Raptors

Osprey

Black-shouldered Kite
Brown Goshawk
Collared Sparrowhawk
Wedge-tailed Eagle
Little Eagle
Spotted Harrier
Swamp Harrier
Peregrine Falcon
Spotted Owl
Barn Owl
Kookaburra

Aerial

Tawny Frogmouth
Owlet-nightjar
Spotted Nightjar
Rainbow Bee-eater
White-backed Swallow
Tree Martin
Fairy Martin
Little Woodswallow

Pandion haliaetus
Elanus notatus
Accipiter fasciatus
Accipiter cirrhocephalus
Aquila audax
Hieraaetus morphnoides
Circus assimilis
Circus aeruginosus
Falco peregrinus
Ninox novaeseelandiae
Tyto alba

Dacelo novaeguineae

Podargus strigoides
Aegotheles cristatus
Caprimulgus guttatus
Merops ornatus
Cheramoeca leucosternum
Cecropis nigricans
Cecropis ariel
Artamus minor

Emu

Spotted Bowerbird
Pied Butcherbird
Grey Currawong
Torresian Crow

Stubble Quail
Painted Button-quail
Laughing Dove
Diamond Dove
Common Bronzewing
Crested Pigeon
White-tailed Black
Cockatoo
Regent Parrot
Budgerygah
Red-capped Parrot
Western Rosella
Many-coloured Parrot
Blue Bonnet
Bourke’s Parrot
Elegant Parrot
Painted Firetail

Dromaius novaehollandiae
Chlamydera maculata
Cracticus nigrogularis
Strepera versicolor
Corvus orru

Coturnix pectoralis
Turnix varia
Streptopelia senegalensis
Geopelia cuneata
Phaps chalcoptera
Ocyphaps lophotes
Calyptorhynchus baudinii

Polytelis anthopeplus
Melopsittacus undulatus
Purpureicephalus spurius
Platycercus icterotis
Psephotus varius
Northiella haematogaster
Neophema bourkii
Neophema elegans
Emblema picta

Graminivores

insectivores

Ground

Bush Stone-curlew
Pallid Cuckoo
Black-eared Cuckoo
Rufous-tailed Bronze-
cuckoo

Shining Bronze-cuckoo
Southern Scrub-robin
Scarlet Robin
Hooded Robin
White-breasted Robin
Western Yellow Robin
Jacky Winter
Crested Shrike-tit
Gilbert Whistler
Golden Whistler
Rufous Whistler
Crested Bellbird
Restless Flycatcher
Cinnamon Quail-thrush
Nullarbor Quail-thrush

insectivores

Burhinus magnirostris
Cuculus pallidus
Chrysococcyx osculans
Chrysococcyx basalis

Chrysococcyx lucidus
Drymodes brunneopygia
Petroica multicolor
Melanodryas cucullata
Eopsaltria georgiana
Eopsaltria griseogularis
Microeca leucophaea
Falcunculus frontatus
Pachycephala inornata
Pachycephala pectoralis
Pachycephala rufiventris
Oreoica gutturalis
Myiagra inquieta
Cinclosoma cinnamomeum
Cinclosoma alisteri

Nectarivores

Purple-crowned Lorikeet

Little Wattlebird
Spiny-cheeked Honeyeater
Singing Honeyeater
Purple-gaped Honeyeater
Yellow-plumed Honeyeater
Grey-fronted Honeyeater
White-plumed Honeyeater
Brown-headed Honeyeater
White-naped Honeyeater
Brown Honeyeater
New Holland Honeyeater

White-cheeked Honeyeater
White-fronted Honeyeater
Tawny-crowned
Honeyeater
Western Spinebill

Pied Honeyeater

Glossopsitta

porphyrocephala
Anthochaera chrysoptera
Acantliagenys rufogulans
Lichenostomus virescens
Lichenostomus cratitius
Lichenostomus ornatus
Lichenostomus plumulus
Lichenostomus penicillatus
Melitlireptus brevirostris
Melithreptus lunatus
Lichmera indistincta
Phylidonyris

novaehollandiae
Phylidonyris nigra
Phylidonyris albifrons
Phylidonyris melanops

Acanthorhynchus
superciliosus
Certhionyx variegatus

63

Journal of the Royal Society of Western Australia, Vol. 62, Parts 1-4, 1979.

Waterbirds

Shore and sea birds

Great Crested Grebe
Hoary-headed Grebe
Darter

Little Pied Cormorant
Black Cormorant
Little Black Cormorant
White-necked Heron
Large Egret
Nankeen Night-heron
Freckled Duck
White-eyed Duck
Musk Duck

Black-tailed Native-hen
Dusky Moorhen
Swamphen

Podiceps cristatus
Poliocephalus poliocephalus
Anhinga melanogaster
Phalacrocorax melanoleucos
Phalacrocorax carbo
Phalacrocorax sulcirostris
Ardea pacifica
Egretta alba
Nycticorax caledonicus
Stictonetta naevosa
Aythya australis
Biziura lobata
Gallinula ventralis
Gallinula tenebrosa
Porphyrio porphyrio

Little Penguin
Australian Pelican
Black-faced Cormorant
Pied Oystercatcher
Hooded Dotterel
Black-fronted Dotterel
Silver Gull
Caspian Tern
Roseate Tern
Bridled Tern
Crested Tern

Eudyptula minor
Pelecanus conspicillatus
Phalacrocorax fuscescens
Haematopus ostralegus
Charadrius rubricollis
Charadrius melanops
Larus novaehollandiae
Hydroprogne caspia
Sterna dougalii
Sterna anaethetus
Sterna bergii

04

Journal of the Royal Society of Western Australia, Vol. 62, Parts 1-4, 1979, p. 65-74.

Aspects of Science in Western Australia 1829-1979.

A volume to commemorate the 150th Anniversary of the founding of Western Australia.

Niche theory and macropodid nutrition

by J. E. Kinnear and A. R. Main

Department of Fisheries and Wildlife, Wildlife Research Centre, P.O. Box 51, Wanneroo, W.A. 6056
(formerly Zoology Department, University of Western Australia)

Zoology Department, University of Western Australia, Nedlands, W.A. 6009

Abstract

Arid zones are universally acknowledged to be nutritionally hostile environments;
this paper sets out to explain the observed capacities of the “ruminant-like” macro-
podid marsupials to survive in such environments by relating their nutritional
biology to the concept of the niche. New insights are achieved by this approach.
The nutritional niche is defined as a subset of the Hutchinson niche space;
it is shown that the symbiotic interaction between the pregastric microbes and
the host mammal enables the host to occupy an expanded nutritional niche.
It is predicted that under specific and well-defined conditions, mammals with
pregastric fermentation (PGF — i.e. ruminant and ruminant-like species) are
competitively inferior to non-PGF herbivores in a nutritional sense, but they avoid
the consequences of this competition by occupying expanded nutritional niches
which exclude competitors. This nutritional theme of niche expansion common to
all mammals with PGF is shown to be nutritionally flexible allowing specialization
in the nutritional niche which is more pronounced than external morphology

would indicate.

Introduction

Humans in arid conditions are impressed by
the heat, shortage of water, and absence of
shade, and it is natural to believe that arid-
zone animals occur where they do because they
can handle these stresses. We therefore tend
to concentrate our studies on measuring the
performance of arid-zone animals under labora-
tory conditions which are believed to represent
in some way field conditions in arid places.

These laboratory studies measure achievement
of animals, i.e., what the animal can endure,
because when we set up laboratory experiments
we take care that the animal cannot behaviour-
ally avoid the conditions we wish to impose on
it. The results of these studies are thus
measures of survival, but frequently we have no
measure of whether the animal experiences such
conditions in the field or not.

However, in order to persist, a species must
not only endure environmental stresses but also
maintain a population, and this is done only
when offspring survive and grow to reproductive
age. Even this is not sufficient to ensure per-
sistence, because an organism also has to con-
tend with competitors for resources which may
be in short supply. Thus three stages are
involved: (a) Survival, in which physiological

performance may be important but behavioural
avoidance may be equally important; (b)
Growth, in which nutrition, especially dietary
protein, vitamins, essential lipids, minerals and
energy, are important; (c) Reproductive success

(also dependent on adequate nutrition), which
ensures that aging animals are replaced by off-
spring. These stages are not equally easy to
analyse. The first has fewer difficulties and is
commonly studied as environmental physiology
or ecophysiology. It is generally assumed that
the present capacities of animals to maintain
life in the face of hot arid conditions represent
the culmination of all the selection there has
been over the period that has elapsed since the
environment became arid.

It is now thought that the onset of aridity
occurred much earlier than formerly believed.
It appears that a trend towards aridity started
in Late Miocene times (Bowler 1976), and was
accentuated by climatic oscillations of wet and
dry periods in the Quaternary. Associated with
these oscillations there is a well-documented
extinction of many large marsupials and in those
cases where extinction did not occur, selection
for smaller-sized species occurred (Bartholomai
1972, 1975; Marshall 1973); but even the smaller
forms have continued to disappear as, for
example, potoroos were only taken shortly after
the settlement of Western Australia and not
subsequently. Thus it would appear that
although a number of species have become rare
or extinct because of European settlement,
evidence from fossil and subfossil deposits
implies that at the time of the colonisation of
Western Australia, the faunal assemblage
represented a terminal stage of a long process
of extinction and a reduction of faunal diversity
which began much earlier.

82710 ( 5 )

65

Journal of the Royal Society of Western Australia, Vol. 62, Parts 1-4, 1979.

It is a common observation that when faunal
diversity is reduced, the surviving species
frequently expand their range or occupy habitats
in which they were formerly never found;
ecologists describe such situations as competitive
release or expansion of the realised niche. For
example, the habitat occupied by the quokka
Setonix brachyurus on Rottnest Island, and the
tammar wallaby Macropus eugenii on the
Abrolhos, is quite unlike the preferred situation
on the mainland; both cases seem to indicate
habitat expansion made possible in part by the
absence of competitors. Another example of
ecological release from constraining environ-
mental variables has been observed in the case
of kangaroos, as when pastoral water and
improved grazing conditions enable kangaroos
to increase in numbers (Ealey 1962, 1967;

Newsome 1975). But these are exceptions
because, in general, there is little evidence to
suggest that range expansion has occurred in
the case of the surviving fauna, and on this
basis we can infer that the present depauperate
fauna exists in competitive equilibrium for
environmental resources.

We are still left with the question as to why
so many species became extinct. And can we,
by studying the ability of the faunal elements
which survived the extinction, make any state-
ment as to why the current fauna persisted?
Was it physiological capacity, reproductive
efficiency, nutritional flexibility or behavioural
avoidance?

Some of these aspects have been previously
dealt with by Main (1968, 1970, 1976, 1978) and
Main and Bakker (in press). Here we propose
to deal with the physiology of nutrition insofar
as this has relevance to the trend to aridity
already discussed. We will argue that the
dominant herbivorous marsupials — the macropo-
dids, i.e., kangaroos, wallabies, rat kangaroos —
have evolved a flexible system of alimentation
appropriate to the arid zone which has a
recognisable nutritional theme. Moreover, we
will attempt to show that this flexible nutritional
theme has many variations which have been
moulded by competitive interactions as predicted
by ecological concepts and the theory of the
niche. Finally, we will also attempt to show
that the nutritional adaptations displayed by
the macropodids have compromised their
energetic and reproductive efficiencies.

Accordingly, this paper will not be a literature
review of the physiology of desert animals as
this subject has been more than adequately
covered by recent reviews (Brown 1974, Russell
1974). Instead we propose to introduce a new
approach to the nutritional physiology of the
macropodid marsupials and then to discuss some
of the biological implications of this approach.
We will begin our account by introducing some
basic concepts of niche theory.

The niche: an abstractly inhabited hypervolume

Niche is a term that was first used to indicate
the role or function of a species in the com-
munity. The concept was not well defined, and
as a result usage was not precise and the term
became almost meaningless. In 1957, the con-

cept of niche acquired elements of mathematical
rigour and precision previously unrealised, due
to the efforts of G. E. Hutchinson (1957) who
conceived of a new way of defining the niche.

In set-theoretic terms, the fundamental niche
(N) of a given species is a n-dimensional hyper-
volume or space. Hutchinson considered all of
the environmental and biological factors (not
ncessarily independent) that affect the survival
of a species, and to every variable or factor he
assigned an axis. On each axis he specified
arbitrary limits, the limits denoting a region
within which a given species can survive and
persist.

To illustrate Hutchinson’s concept, we will
consider two familiar variables that affect the
survival of desert herbivores — temperature and
available water in plant tissues. A given species
will survive and reproduce within a certain
range of temperature and water concentration
in plant tissues. If these ranges are linearly
ordered on a pair of rectangular co-ordinate axes
(Fig. 1A), a region (in 2 space) is defined within
which the species can persist. In effect we have
specified the physiological tolerances of a given
species with respect to temperature and available
plant water. The concept is generalised by
considering all relevant variables, i.e., a third
axis would define a volume or 3 space; n-axes
would define a hypervolume of n-space and
thus a species’ fundamental niche.

Upon defining the fundamental niche,
Hutchinson proceeded to qualify the concept by
considering biological interactions (e.g. com-
petition) among organisms. He reasoned that
because of biological interactions, a species need
not exist within the full extent of its funda-
mental niche; this contracted niche space,
Hutchinson termed the realised niche.

An assumption, indeed a biological axiom,
states that at equilibrium, no two sympatric
species can occupy the same niche space. This
assumption is based on the competitive-exclusion
principle (see Hardin 1960) ; we will have
occasion to invoke this principle as some impor-
tant predictions flow from it.

On considering the foregoing, a little reflection
reveals that much of the physiological literature
represents efforts by physiologists to define a
species’ fundamental niche. Any studies that
characterise a species’ tolerance or requirement
for an environmental factor can be interpreted
as characterising axes or subspaces of a species’
fundamental niche. Furthermore, if a physio-
logical adaptation increases the tolerance of an
organism to a stressful factor, or lowers the
requirement for an environmental variable, then
such adaptations may be defined as expanding
the fundamental niche. We suggest that this
is a useful way of describing the biological
significance of physiological adaptations.

The merits of this approach will be particu-
larly apparent when we discuss the biological
implications of macropodid nutrition, as some
new insights have been gained by applying the
above ecological concepts. However, before we
do this it will be necessary to describe some
background material.

66

Journal of the Royal Society of Western Australia, Vol. 62, Parts 1-4, 1979.

Early studies on marsupial nutrition

Pioneering studies on macropodid nutrition
began when H. Waring approached R. Moir and
M. Somers of the Institute of Agriculture about
the nutritional significance of the unusually
large forestomach of the Rottnest quokka
( Setonix brachyurus) . Their studies revealed
that the quokka was “ruminant-like”. Bacteria
were present in huge numbers in the stomach
contents and microbial end-products — volatile
fatty acids (VFA) — were found in concentra-
tions comparable to those in domestic ruminants
(Moir et al. 1956). Subsequent work has amply
confirmed their original conclusions, and the
ruminant-like physiological convergence has
since been extended to other macropodids as
well as some eutherians (Moir 1968).

Additional nutritional studies were stimulated
by Ealey’s (1962, 1967) investigations on the
euro or hill kangaroo (Macropus robustus) in
the Pilbara. This work was initiated in response
to the demands of pastoralists to control the
euro which was viewed by the industry as a
serious economic pest. One of the results of
Ealey’s research was the finding that pastoral
watering points had enabled the euro population
to increase. Under natural conditions, the euro
population is limited by intraspecific competition
for heat refuges in the form of caves in break-
away country — a necessary requirement to main-
tain water balance. By making free water
readily available, the pastoral industry had
removed a limiting environmental factor of the
niche space of the euro; water balance could
be maintained in the absence of heat refuges.

It is significant that the euro population had
expanded its realised niche by interacting with
a dimension (water) of an agro-ecosystem. Thus
the euro is not adapted to the extreme heat as
for example are some African gazelles (Taylor
1972) which have expanded their fundamental
niches in a physiological way. Survival of the
euro under natural conditions is achieved by
appropriate behavioural responses in suitable
habitats — a tactic which many animals of the
arid zone employ.

While there appeared to be no unusual physio-
logical adaptations by the euro to water, studies
of their dietary preferences implicated physio-
logy (Storr 1968). Euros were maintaining their
numbers on an extremely fibrous low-protein
food source (e.g. Triodia sp.) which did not
support domestic sheep. How the euro popu-
lation was adapted to such a low-grade nutri-
tional environment remained unknown until G.
Brown investigated the problem. His findings
were quite remarkable, and his thesis represents
a study that was the first of its kind in the
sense that no-one had ever determined the
nitrogen requirement of a wild species.

The remarkable results obtained from nitro-
gen-balance trials in the euro revealed the
minimum nitrogen requirement of the euro to
be 50% lower than that of sheep. The results
were fully consistent with the field observations
on the diet of the euro, i.e., the euro is physio-
logically adapted to low protein diets; it has
expanded its niche space by having a low

requirement for environmental nitrogen (Brown
1964, Brown and Main 1967, Brown 1969).

Perhaps equally as significant was Brown’s
explanation for the low nitrogen requirement
of the euro. He correctly hypothesised that the
euro must have a lower standard metabolism,
and that the euro recycled urinary nitrogen.
Two types of niche expansion are therefore
responsible for the lower nitrogen requirement —
the first is an expansion of the fundamental
niche made possible by a lower rate of metabo-
lism; the second is an interaction of the animal
with its stomach micro-organisms, but this
represents an expansion of the realised niche.
The distinction is a subtle one which has not
been recognised, nor have the consequences been
appreciated.

Niche theory and macropodid nutrition

The arid zone is not only noted for its heat
and water stresses, but also is characterised by
cyclic and unpredictable bursts of primary pro-
duction followed by long periods of low produc-
tivity. During these unproductive periods the
standing crop may still be extensive, but the
vegetation becomes nutritionally hostile, for one
routinely observes in the case of macropods, a
loss of body condition (Kinnear and Main 1975)
and sometimes heavy mortality in the population
(Ealey 1962, Main 1970).

Herbivorous mammals have never directly
solved the nutritional challenge of the arid
zone. They survive and persist because they
have entered into a complex biological association
(symbiosis) with microbes. It is significant that
they have largely adapted their physiology to
maintain the association and to exploit its
benefits.

The associations follow two basic patterns :
microbes are housed and continuously cultured
in two distinct regions of the alimentary tract —
either the hindgut (i.e., post-ileo) or both hind-
gut and foregut. In the latter case, the foregut
population is invariably dominant in terms of
microbial numbers and gut volume. This design
of housing microbial populations in the foregut
has been described as pregastric fermentation
(PGF) (Moir 1965, 1968), a useful term which
rigorously distinguishes PGF mammals from
those species that house microbial populations
solely in the hindgut (e.g. lagomorphs, perisso-
dactyls) .

PGF mammals are represented principally by
ruminants and, in the case of marsupials, by
the macropodids which are also referred to as
being “ruminant-like”. Both groups underwent
an explosive radiation during middle/late Terti-
ary times. These evolutionary events, which
occurred independently, are correlated with
climatic changes that caused seasonal or discon-
tinuous periods in primary production. It is
thus a common nutritional solution, and a
global phenomenon, with many variations on a
theme.

The biology of this nutritional theme of hous-
ing a microbial fermentation system in the fore-
gut has in our view never been adequately

67

Journal of the Royal Society of Western Australia, Vol. 62, Parts 1-4, 1979.

explained nor the purpose understood. This is
so despite intensive research by nutritional
physiologists and animal scientists; it is gener-
ally believed that PGF is primarily an adapta-
tion which enables the host animal to exploit
fibrous (cellulosic) plant structures as a source
of energy.

The rumen and its microbes are also acknow-
ledged to have the properties of an ecosystem.
Some attempts have been made to analyse the
system as such, but these attempts have not
yielded any fresh insights. However, new
insights and properties emerge when this bio-
logical association is interpreted in terms of
niche theory.

The nutritional niche

The generality of Hutchinson’s fundamental
niche concept allows one to partition the funda-
mental niche into subsets of axes. We define
the fundamental nutritional niche as a subset of
axes restricted to those axes representing
nutrient variables (e.g., essential amino acids,
carbon, phosphorus, essential lipids, vitamins,
H 2 0, etc.) essential to the persistence of a
species.

It follows from the above that, if a species is
to occupy a niche, there has to be a correspon-
dence between a species’ nutrient requirement
(NR) and its nutritional niche space (NNS).
In abstract terms this means a species must be
able to map its NR set into an NNS set. If a
species cannot perform this mapping then in
concrete terms its nutrient source is lacking in
nutrition in one or more ways; persistence is
therefore not possible.

Competition, niche and PGF

A species may be capable of mapping its
nutritional requirements into a nutritional niche
space, but it can be prevented from occupying
a niche space by more efficient competitors.
This leads us to examine the competitive abilities
of mammals with PGF; we will first consider
the theoretical aspects and then we will review
the available evidence.

The events that occur in the pregastric diver-
ticulae are usually referred to as “digestion”,
but this is clearly an inappropriate word; a more
accurate description of the processes that occur
is decomposition. Food of PGF mammals is not
initially digested, but is partially decomposed
(to varying degrees) by microbes; associated
with these events are the growth and mainten-
ance of a microbial population — the pregastric
culture.

One of the consequences of microbial decom-
position is that some of the dietary components
are rendered digestible (e.g. fibre) while other
components can be improved in nutritive value
(e.g. the essential amino-N and vitamin con-
tent) ; on the debit side, decomposition, as the
term implies, leads to energy losses and mineral-
isation of the host’s food. It is for these

reasons that we observe the presence of the
decomposition products — NH 4 , H 2 S, CHi and C0 2
— in the stomach diverticulae.

To emphasise the above, we can say that
ruminants and their analogues do not initially
digest their food — they rot it, or at least a
major part of it. In order to achieve this, these
mammals have domesticated microbes and as
a consequence they have acquired a unique
trophic level (Kinnear and Main 1975).

In acquiring a microbial trophic level PGF
mammals have increased the links in their food
chain. We recognise: plants -> bacteria ->
protozoa (if present) mammal. If this were
all we knew about these animals, we could still
confidently predict on the basis of trophic
energetics (and thermodynamics) that macro-
podids, ruminants, etc., could not possibly be as
productive as non-ruminant animals. This is to
be expected because productivity calculations
must reckon with an additional trophic level,
and the inevitable losses associated with the
energy and material transfer between trophic
levels. The losses are not as great as that
observed in more conventional food chains
because of the anaerobic (fermentative) prop-
erties of the microbial trophic level; nonetheless,
as we will show, the losses are still appreciable.

Our trophic analysis has led us to predict that
ruminants and their analogues are comparatively
inefficient at transforming food into animal
biomass. This conclusion is consistent with the
experimental evidence — indeed it is an acknow-
ledged fact in the field of animal production
(Reid 1970). For example, Armstrong and
Blaxter (1961) demonstrated that carbohydrate
yields 11 to 30% more energy for production
when digested post-ruminally than when decom-
posed in the rumen. Similar results occur with
regard to protein utilization by ruminants. In
reviewing this aspect, Smith (1975) concludes:
“It seems that the degradation of food protein
in the rumen and the use of the products for
the synthesis of microbial protein (even though
this may be of high biological value) is a very
inefficient and wasteful means of using protein
resources.” In recognition of these inefficiencies
it is not surprising therefore to note that animal
scientists are endeavouring to develop feeding
regimens which enable the rumen microbes to
be bypassed (Owens and Isaacson 1977).

To summarise our discussion to this point, we
have shown that PGF mammals are relatively
inefficient in converting food resources into
animal biomass. But this conclusion makes one
uneasy for it seems to create a paradox. Given
that PGF mammals suffer from measurable
inefficiencies, how can we explain the success of
the group? What has happened must be this:
these animals have made energetic and nutri-
tional compromises in exchange for the ability
to exploit and colonise niches that nutritionally
exclude most other mammals with different
alimentary systems; in other words, they do not
compete with non-PGF herbivores for the same
sort of food.

68

Journal of the Royal Society of Western Australia, Vol. 62, Parts 1-4, 1979.

The nutritional niche space of mammals
with PGF

Since we have reached the conclusion that
PGF mammals cannot compete with non-PGF
herbivores, we have raised the question — What
is their nutritional niche?

Now the symbiosis between the microbial
trophic level and the host animal is a biological
interaction, and according to Hutchinson (1957)
biological interactions affect the dimensions of
the niche hypervolume. Competition is known
to contract the niche hypervolume while the
converse — symbiosis, which is really negative
competition — should therefore expand the niche
space. Thus mammals with PGF should be
capable of occupying expanded niches. Again
the literature supports our predictions — consider
the following evidence.

Nutritional niche expansion means that an
organism can lower its requirement for a
nutrient or increase its tolerance to excess of a
nutrient or both. There are many examples of
this in the ruminant literature, and perhaps the
most impressive examples are the demonstra-
tions which show that domestic ruminants can
be maintained in a healthy and productive state
when fed diets containing non-protein nitrogen
(NPN) as a sole replacement for protein N.
Virtanen’s (1967) long-term feeding trials with
high-yielding milk cows are outstanding
examples of a ruminant’s ability to eat foods
deficient in essential nutrients, i.e. essential
amino acids, water-soluble vitamins and
essential lipids. With regard to other ruminant-
like mammals little work has been done, but
Brown (1964, 1969) replaced part of the protein
N with urea N (30% of N intake) and kangaroos
maintained N-balance. The evidence for the
recycling of N by camels (Schmidt-Nielsen et al.
1957) and by wallabies (Kinnear and Main 1975)
is indicative of NPN utilisation. Recycling of N,
of course, is another example of niche expansion
— it lowers the environmental nitrogen require-
ments of an animal.

We have so far focused our discussion on
essential nutrients, pointing out that the pregas-
tric microbial/mammal symbiosis has resulted
in niche expansion by eliminating the need for
an array of essential nutrients. But mammals
with PGF can also use microbes to detoxify
plant poisons (e.g. oxalates, alkaloids; Freeland
and Janzen 1974), which also results in niche
expansion because the microbes have extended
the host’s tolerances to toxic substances in the
food.

It should be stressed that the foregut sym-
biosis has expanded the realised nutritional
niche, and has not affected the host’s funda-
mental nutritional niche. Ruminants still
require a full array of preformed essential
nutrients; without their pregastric microbes
they would require a balanced diet complete in
all respects.

Evidence for nutritional niche expansion

Nutritional niche expansion means that under
natural conditions, PGF mammals should be
eating foods that are unnutritious in one or

more ways. The evidence supporting this pre-
diction is sparse for the simple reason that
dietary studies on wild herbivores have been
limited to what an animal eats, and occasion-
ally to the gross chemical composition of the
diet, e.g. nitrogen, lipids, fibre, minerals, etc.
What is necessary to prove our predictions are
dietary analyses which measure the essential
amino acids, vitamins, essential lipids and toxic
substances in an animal’s natural diet.

Unfortunately, these analyses are difficult to
perform, for they require sophisticated equip-
ment and technical expertise; furthermore, there
are also technical difficulties in the field because
it is difficult in most instances to define what
plants a herbivore i s eating, the amounts con-
sumed, as well as the digestibility. In view of
this situation, what seemed obviously necessary
to us was to find a ruminant-like animal with
simple well-defined dietary habits. We were
fortunate in finding such an animal in the
woylie or brush-tailed rat kangaroo (Bettongia
penicillata ) .

In a study of the ecology of woylies near
Manjimup, Western Australia, P. Christensen
(1977) discovered that woylies dig up and eat
small pebble-sized fungi. We also discovered
from stomach content analyses that woylies
were eating a gum exudate from the trunk of
Hakea sp. There was no evidence that woylies
were eating green plants or fibrous materials of
any kind (Kinnear et al. in prep.). Analysis of
these dietary items revealed that the gum con-
tained no detectable nitrogen and that the fungi
were deficient in the essential amino acid (EAA)
lysine, but in addition, the EAA methionine was
in excess. Hence, not only do we find that
woylies are eating a lysine-deficient protein
source, but also we find that the protein source
has an amino acid imbalance. By known stand-
ards of mammalian nutrition woylies, as pre-
dicted, are eating unnutritious foods but are
meeting their nutritional requirements as the
result of microbial synthesis.

Competitive inadequacies of PGF mammals

In nature, niche theory predicts that we should
observe mammals with PGF occupying niches
that exclude other herbivorous mammals with
different alimentary systems. But they also will
be excluded from nutritional niches — in parti-
cular, those niches which provide nutritionally
balanced foods. This does not mean that
mammals with PGF will not thrive on nutritious
diets for they will, but because of their trophic
status, it means they will utilise such foods
inefficiently relative to non-PGF competitors and
thus they will be excluded from such niches.

There are, however, conceivable circumstances
where dietary analyses may reveal that the
foods eaten by some PGF mammals are balanced
and nutritious. We have so far conceived of
four situations:

(1) The food may be nutritionally complete, but
contains toxic substances that are subject to
microbial degradation.

(2) Nutritious foods are not limiting, i.e. a
surplus exists.

69

Journal of the Royal Society of Western Australia, Vol. 62, Parts 1-4, 1979.

(3) PGF mammals may occupy nutritionally
complete niches during seasonal periods, i.e.
the growing season.

(4) Competitors may be absent.

To illustrate the niche space of PGF mammals,
both in the absence and presence of competitors,

we have presented a series of figures represent-
ing the nutritional niche space for two dimen-
sions only. In the absence of competition we
observe simple niche expansion; in the presence
of competitors, PGF animals are excluded from
an area of the two-dimensional niche space (Fig.
1 ).

(A)

er ^

er \>

N.

(B)

en

EN

en

EN 2

en

r

1

en

2

I J

en

(C)

EN

en

(D)

EN

Figure 1. — The niche space restricted to two dimensions for the purpose of illustration (the concept is
generalised to n-dimensions) . A. — Hutchinson’s fundamental niche: factors or variables affecting a given
species are linearly ordered on a pair of rectangular axes and the limits X”^ — X’^ and X”^ — X’^ define

an area (a 2-space) within which a species can survive and persist. B. — The fundamental nutritional niche:
two essential nutrients are linearly ordered and the limits (en” , en’ etc.) denote an area within which a

i i

species survives and persists. C. — The nutritional niche space of a PGF species; the symbiotic interaction
results in simple niche expansion in the absence of non-PGF competitors. The inner area represents the
fundamental nutritional niche. D. — The nutritional niche space of PGF species under competition with non-
PGF species. PGF species are excluded from regions of their fundamental niche space because they are com-
petitively inferior.

70

Journal of the Royal Society of Western Australia, Vol. 62, Parts 1-4, 1979.

Competitive interactions between PGF mammals

An immense number of feeding trials involving
domestic ruminants has revealed that these
animals possess an extremely versatile and
adaptable alimentary system, for they have been
fed an incredible array of foodstuffs — from
newsprint to anchovies. These animals seem
capable of eating almost anything, anywhere;
their system of alimentation would seem to be
the ultimate nutritional solution.

But such impressions are tempered when
viewed from an ecological perspective, for eco-
logical theory demands that co-existing species
must avoid excessive niche overlap. We should
therefore expect and observe niche partitioning
of food resources by sympatric species of
mammals with PGF. Furthermore, as a corol-
lary, we should expect to observe specialisations
in physiology and morphology.

Recent studies have borne out these ecological
predictions. Hofmann (1973) has produced an
impressive monograph on the African ruminants
which describes, in great detail, the different
morphological adaptations of the rumen which
are correlated with dietary preferences.
Hofmann has related stomach structure to the
type of food eaten, and he has arranged a large
number of African species into three major
groups; (a) concentrate feeders — low fibre/
succulent diets; (b) roughage feeders — fibrous
diets; and (c) concentrate/roughage feeders —
i.e. species that eat either foods depending on
whatever is seasonally available.

Less information on nutritional-niche parti-
tioning exists for the ruminant-like macropodids
but from the available information a similar
pattern is emerging. For example, the woylie
rat-kangaroo stomach differs substantially from
that of wallabies and kangaroos; moreover it is
a concentrate feeder (as is the Tasmanian
potoroo — Guiler 1971) which ensures that it
does not overlap or compete with the three
species of macropods using the same habitat
(Kinnear et al. in prep.).

65

55

o

35

25 i i i i i ,

10 20 30 40 50 60 70

DRY MATTER INTAKE

Figure 2. — The performance of 3 species of kangaroo
in digesting fibrous diets. As dry matter intakes
(dry matter wt. kg°-75) increase, the percentage of
dry matter digested (DMI (%) ) tends to decline
slightly for all species, but the significant points
are the differences between the 3 species in ex-
tracting digestible energy from fibrous diets (after
Prince 1976 — with permission).

With respect to physiological adaptation and
specialisation, differences between the three
common species of kangaroos have been docu-
mented by Prince (1976). In comparative feed-
ing trials, a standard diet containing fibre was
fed to western grey kangaroos (Macropus fuli-
jinosus) , red kangaroos (Megaleia rufa) and
euros; appreciable differences in digestive effici-
encies were noted.

The differences are illustrated in Figure 2
where it is evident that the three species differ
significantly in their abilities to extract digest-
ible energy from fibrous diets. The red kan-
garoo is least efficient in obtaining energy from
fibre, which means that it must seek out
nutritional niches that provide low-fibre foods;
in the wild, they are known to do this (Newsome
1975). The opposite extreme is the grey kan-
garoo which is the most efficient, and Prince
points out that the grey, which has a southern
distribution, is more subject to cold stress (due
to low temperatures and winter rains), and
hence the ability to obtain more energy from
fibre is an adaptive specialisation to the environ-
ment and probably necessary to assure reproduc-
tive success. Finally, the euro which is

predominantly northern in its distribution
(summer rainfall) must also contend with fibre
but it can afford to be less efficient.

The prevailing view held by most animal
scientists is that the rumen has evolved to cope
with fibrous foods. But it is now clear that this
view is untenable because PGF mammals eat a
wide array of plant materials, many of which
are low in fibre. Hofmann’s study shows that
the fibre-consuming ruminants are merely
specialists and we also observe the same sort of
specialisation in the case of macropods, i.e. the
grey kangaroo and the euro.

Niche theory provides a more general explan-
ation: the function and purpose of PGF is to
allow these animals to occupy expanded nutri-
tional niches. At this stage (apart from our
woylie study) we cannot support this explana-
tion by factual data because the need for such
data has not been recognised, and hence it has
not been systematically collected. We hope that
this challenge will be taken up.

Discussion

There seems to be little reason to doubt that
pregastric fermentation is indeed a flexible
system of alimentation that has been moulded
by the pressures of natural selection and com-
petition. We observe a plethora of species (ca.
50 species of marsupials and > 200 eutherian)
frequently existing sympatrically in closely-knit
associations that seem to defy the competitive
exclusion principle. Mammals with PGF are the
dominant vertebrate grazer/browsers through-
out the world in contrast to the hindgut-fer-
menting mammals which are relict survivors of
a previously dominant group (Moir 1965).

The climatic factors associated with the rise
of PGF mammals are cold temperatures and
aridity which do not favour continuous plant
growth. This forced mammals to satisfy their
complex nutritional requirements from vegeta-

71

Journal of the Royal Society of Western Australia, Vol. 62, Parts 1-4, 1979.

tion that was either in a state of dormancy,
senescent, or dead. The PGF mammals solved
this nutritional challenge in a peculiar way —
they largely abandoned herbivory by acquiring
a decomposer trophic level which they housed
in their forestomachs.

It is this microbial trophic level which imparts
nutritional flexibility and which also enables
nutritional specialisation. The key to the success
of PGF mammals has been the near abandon-
ment of herbivory in favour of linking their
food chain to a captive-decomposer trophic level.
A significant feature of this trophic level is its
affinities with the soil-decomposer ecosystem
(Hungate 1960), for such comparisons indicate
that PGF mammals have domesticated anaerobic
soil micro-organisms which have retained their
ecological function — that of decomposing organic
matter. When these relationships are recog-
nised and appreciated, it is easy to understand
why macropods of the arid zone can survive for
extended periods while consuming drought-
affected vegetation that is nutritionally equiva-
lent to plant litter.

However, this nutritional theme of housing a
microbial trophic level in the foregut is not
without its shortcomings for there are com-
petitive and energetic disadvantages. PGF
mammals are inherently less efficient in con-
verting and transforming food into animal bio-
mass and, moreover, this inefficiency would seem
to have reproductive consequences.

If one surveys the mammalian literature (e.g.
Walker 1975) for data on fecundity, a pattern
emerges: eutherian mammals with PGF usually
bear 1-2 young and this is so regardless of size;
macropodid marsupials are uniformly consistent
in that they give birth to only one young at a
time. Thus it would appear that mammals with
PGF have difficulty in allocating their food
resources to the production of young; this
characteristic is in sharp contrast to the repro-
ductive performance of some non-PGF herbi-
vores, e.g. the rabbit and many rodents.

The macropodid marsupials are further
handicapped energetically for reasons that they
have a lower metabolic rate (MR) relative to
most eutherians (Dawson and Hulbert 1969,
Kinnear and Shield 1975). Dawson (1973) has
advanced the view that marsupials are ener-
getically primitive, but such views do not help
to explain why so many eutherians find it
advantageous to generate metabolic power and
heat in excess of their homeothermic require-
ments.

Kinnear and Shield (1975) suggest some
advantages: “A high metabolic rate enables
rapid synthesis and storage of energy as well as
accelerated neonatal development and growth
during favourable periods. All of these would be
advantages to an animal facing future energy
stress.” The authors cite as an example the MR
of the white-tailed deer ( Odocoileus virginianus)
which is up to twice as high in summer than
winter. This reduction in MR allows for a
slower rate of utilisation of stored energy
reserves which are rapidly accumulated during
peaks of primary production during summer.

Metabolism is an adaptive physiological para-
meter which not only provides heat to maintain
a mammal’s body temperature but also deter-
mines an animal’s biosynthetic potential, i.e.
the rate at which it can create biomass. With
this point in mind we can appreciate the short-
comings in combining the energetic inefficiencies
associated with PGF with a marsupial rate of
metabolism, and this would seem to have some
relevance to the extinction of the giant macro-
podids. These species did not survive the
climatic oscillations of wet and dry periods in
the Quaternary which were superimposed on an
overall trend of increasing aridity. The periods
of primary production favouring reproduction
and growth would have been too short for
recruitment because these giant and metabolic-
ally sluggish marsupials would have lacked
sufficient biosynthetic power to maintain a
population. It is significant that the surviving
macropodid descendants are smaller-sized
species and, moreover, they are sexually dimor-
phic with the female being smaller and becoming
sexually mature at an earlier age than the male.

Thus, in the case of extant kangaroos, each
species has adults of two sizes, and when several
species occur together as in the Goldfields region
of Western Australia where the red, euro and
grey are found, there are effectively adults of
six size classes. The reason for the contain-
ment of these species within their ranges and
realised niches requires further study, as Main
(1978) has suggested for the western and eastern
grey Kangaroos in south-eastern Australia.
Should such studies reveal, as seems likely, that
habitat specialisation and not inter-species
competition is the regulating factor, then
habitat specialisation and the nutritional niche
of each species are much more highly developed
than external morphology would suggest.

From the above reasoning on habitat and
niche specialisation, it would seem to follow
that habitat alteration and not competition
caused past extinctions, and conversely the
extant or successful species persisted because
they can match their needs even when dietary
resources are of poor quality, and so survive to
reproduce.

We conclude our account on the following
points: within the aerobic zones of the bio-
sphere there are no empty nutritional niches,
for all primary production on balance is
utilised by consumers or decomposers. Extinc-
tion of faunas therefore does not necessarily
create or open up nutritional niches, for such
niches can be partitioned in a different manner
until a new equilibrium is reached between con-
sumers and decomposers. Nutritional niches
disappear with the extinction of plant species
and communities and, conversely, new niches
arise when new plant communities evolve.

It is noteworthy that in arid and semi-arid
Australia the one ungrazed habitat is acacia
scrub extending from a sandy coastal situation
inland to the mulga scrub. No marsupial sur-
vived to exploit this nutritional niche yet, this
habitat with its tall acacia should have been
within the range of the giant marsupials, many

72

Journal of the Royal Society of Western Australia, Vol. 62, Parts 1-4, 1979.

of which are believed to have been browsers.
Thus while the Australian fauna has many
parallels with its eutherian counterparts, it is
significant that the present fauna has no equi-
valents to the large African browsers, e.g. the
Giraffidae. Apparently, the large macropod
browsers were incapable of adapting to aridity
by dwarfing as in other macropods (Main 1978),
because smaller animals are undoubtedly denied
the browse on trees and shrubs.

Elsewhere the eutherian browsers persisted in
the face of increasing aridity, often by using
water holes (Leuthold 1977), and it is signi-
ficant that when such large species were re-
leased in Australia, e.g. the feral ungulates such
as the camel, they have successfully colonised
the acacia habitat.

Acknowledgements. — We thank A. Cockson and M.
Onus for constructing the figures, and acknowledge
financial assistance from C.S.I.R.O., the Australian Re-
search Grants Committee and the University Research
Grants Committee to A.R.M. throughout the course of
these investigations.

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74

Journal of the Royal Society of Western Australia, Vol. 62, Parts 1-4, 1979, p. 75-82.

Aspects of Science in Western Australia 1829-1979.

A volume to commemorate the 150th Anniversary of the founding of Western Australia.

Vegetation mapping in Western Australia

by J. S. Beard

6 Fraser Road, Applecross, W.A. 6153

Abstract

Interest in conservation and wildlife management has increased the value of
vegetation mapping as a form of resource inventory. Western Australia achieved an
early lead in this field with the work of Diels (1906) and Jutson (1914) but thereafter
initiative passed to the eastern States for a number of years until the establishment of
a State-wide vegetation survey in Western Australia in 1964. The aims, objects and
methods of this survey are described and its achievements in publication are listed.

Introduction

There are basically two different ways in which
a field botanist can regard the plant cover of an
area in which he is working. He can see it as
composed of individual plant species, each to be
named and classified, or he can see it in the mass
as vegetation having properties of structure and
life-form. The former is the taxonomist’s view-
point, the latter the biogeographer’s. In the
latter case the existence of species is always of
interest but it is not the prime interest. Some
biogeographers are interested in species as units
of association studying their occurrence and
relative abundance in different habitats, but
others to whom the physiognomy of vegetation
is the dominant interest may regard species as
no more than incidental. Studies of vegetation
are normally accompanied by mapping of vegeta-
tion units so that location and extent as well as
physical characteristics may be documented, and
as techniques of mapping have improved so has
vegetation mapping grown as an art and a
science.

A vegetation map may be defined as one which
shows the nature of the earth’s plant cover.
Sometimes this cover is a natural one, unmodified
by human action, as we still find today in the
unsettled inland areas of the State, and some-
times it has been entirely created by man, as in
intensive agricultural areas. In intermediate
cases, for example, those parts of Western Aus-
tralia that are under pastoral leases, there is a
natural plant cover that has been or is being
modified by man and his domestic animals. A
vegetation map records what is there but in
general crop mapping in agricultural areas is
commonly regarded as a separate field, i.e. land-
use mapping, while a vegetation map in the strict
sense is expected to show natural plant cover
as it exists or used to exist. In settled areas
there is often an attempt to reconstruct a picture
of the original or primaeval plant cover.

The data presented in vegetation maps may
be interpreted in many ways and for many
different purposes, including the following: —

Conservation: Vegetation mapping provides an
inventory of plant communities and plant habi-
tats, which is the basic information required for
planning an adequate system of biological re-
serves. At a later stage a vegetation map of a
flora and/or fauna reserve or national park is
one of the basic necessities for planned manage-
ment and control.

Environmental-impact studies. Facts about
vegetation are a part of necessary basic know-
ledge when assessments of impact upon the
environment made by proposed mining and other
land development are being made.

Potential land use. A vegetation map is a use-
ful basis for more detailed surveys of land poten-
tial for agriculture, pastoral use and forestry,
and for soil and geological surveys. It shows
height and density of vegetation and may be
used as a “going” map for military purposes and
for oil and mineral exploration.

Early work in Western Australia

Until quite recently the art of vegetation map-
ping had been very little practised anywhere in
Australia. Difficulties in travel and communica-
tion, and lack of accurate topographic base maps
were formidable obstacles. Gradually, however,
as interest in conservation and wildlife manage-
ment has grown, interest has grown also in
vegetation mapping as a necessary form of
resource inventory, and several mapping pro-
grammes have been set up, supported by the
Interim Council for the Australian Biological
Resources Study and by State Governments.

It is a common experience in mapping, whether
it be topographic, geological or phytogeographic,
that it begins with some rather imprecise sketch-
maps usually at large scales made by the pioneers

75

Journal of the Royal Society of Western Australia, Vol. 62, Parts 1-4, 1979.

and proceeds by successive revisions to greater
precision at smaller scales. Before the introduc-
tion of aerial photography in the 1930s it was
virtually impossible to map large areas accur-
ately in detail and any precise vegetation map-
ping was confined to local studies. Larger areas
had inevitably to be covered on a sketch-map
basis and involved a large amount of guess-work.
L. Diels for example furnished a coloured sketch-
map at 1:27 000 000 of the whole continent as
the end-papers to his book of 1906 but it cannot
have been intended to give more than a general
impression of the vegetation. Numerous other
small-scale renderings of the vegetation of Aus-
tralia appeared in atlases and geographies
throughout the world, all of them unavoidably
imprecise and unreliable.

It is interesting that in the early period West-
ern Australia was well to the fore. Diel’s book
Die Pflanzenwelt von West- Austr alien (The
plant world of Western Australia) published in
German in 1906, which included the map men-
tioned above, was the only comprehensive and
authoritative account of any Australian vegeta-
tion to be published prior to the first World
War. Western Australia also produced the first
State map of vegetation, when J. T. Jutson in
1914 produced a sketch map at 1:5 000 000 to
accompany his classic work on physiography.
The map was published in colour and distin-
guished nine basic vegetation types, alluding to
two more (fringing forests and mangrove wood-
lands) in the legend. Acknowledgement was
given to the Forests Department as the source of
the information.

In 1928 the Forests Department reprinted the
map with the addition of data contributed by
C. A. Gardner on the distribution of the more
important species of Eucalyptus. This map was
again reprinted in 1967, without revision. In the
meantime Gardner (1942) had published a
general account of the vegetation of the State
which included a small black and white map on
a scale of 1:25 000 000 and this also was still
based on Jutson.

Western Australia’s early lead in this field was
not maintained and the initiative passed in the
1920s to South Australia where the Botany
Department of the University of Adelaide, the
Waite Agricultural Research Institute, the
CSIRO Division of Soils and the State Depart-
ments of Agriculture, Lands and Forests, between
them mapped most of the south-eastern part of
South Australia in terms of vegetation, or of
soils and vegetation, at a considerable range of
scales. This activity came to be known as the
“Adelaide School” and was inspired by Prof. J. E.
Wood of Adelaide who produced an integrated
text and map in 1937. Important contributions
were later made by Blake (1938) in Queensland
and by Beadle (1948) in New South Wales, both
of whom succeeded in mapping very extensive
areas before aerial photography became avail-
able.

The introduction of aerial photography in the
1930s transformed the situation and it became
possible to map vegetation accurately and
rapidly. In Western Australia after World War
II the Forests Department conducted detailed

stock-mapping (1:63 360) of the State Forests in
the south-west, and the pastoral section of the
Department of Lands and Surveys produced a
number of pastoral-classification plans at a scale
of 1:250 000 in which the plant cover was
mapped according to its estimated carrying
capacity. Some other mapping was carried out
by the CSIRO which conveyed vegetation in-
formation without mapping it directly. The
CSIRO Division of Land Research published
three surveys covering almost the whole of the
Kimberley District in the north (Speck et al.
1960, 1964; Stewart et al. 1970) and another
survey (Mabbutt et al. 1963) covered a strip
typical of mulga country in the Murchison region.

These surveys were not vegetation surveys as
such but mapped “land systems”; however, as
vegetation is a component of a land system and
detailed information was given in the explana-
tory text, it was possible to reinterpret for vege-
tation if desired. The same applies to the work
of the CSIRO Division of Soils which has pub-
lished three surveys in the south-west, covering
the Margaret-Blackwood River area (Smith
(1951), the Swan Coastal Plain (McArthur and
Bettenay 1960), and the Merredin area (Bet-
tenay and Hingston 1961). As close correlations
were found between vegetation and soil types
and notes on vegetation were given in the text,
the soil maps could be similarly reinterpreted for
vegetation.

A State-wide vegetation survey

Attempts to depict the vegetation of the whole
of Australia accurately by drawing upon pub-
lished sources had been made by Prescott (1931)
and by Williams (1955) and these highlighted
the serious lack of basic information from
Western Australia which was shown to be by far
the least known of any of the States. In 1964
it was resolved by Prof. M. J. Webb and the writer
to promote a State-wide survey to redress the
situation. This project which was called the
Vegetation Survey of Western Australia has been
carried on since, supported by the Department of
Geography at the University of Western Aus-
tralia where the cartographic work is done, and
in the early stages by the King’s Park Board,
and latterly by the Interim Council for the Aus-
tralian Biological Resources Study, both of which
have provided finance for fieldwork and publica-
tion. Three maps have been contributed by the
Western Australian Department of Agriculture
(Smith 1972, 1973, 1974). Several previous

accounts of the aims, objects and methods of the
survey have been published (Beard 1966a, 1969,
1975a, Beard and Webb, 1974).

The first decision which has to be made on
commencing such a programme is what scale is
to be adopted, and here one has to consider the
area involved, the man-power and finance avail-
able and the duration of the project. Obviously
a survey of Lord Howe Island or even Tasmania
can be made more intensive than one of Western
Australia. It was decided that the 1:1 000 000
scale would be the most generally useful within
the limits of feasibility. It has the advantage of
being the same scale as the International Map of

76

Journal of the Royal Society of Western Australia, Vol. 62, Parts 1-4, 1979.

the World topographic series, and as a French
vegetation map series, the Carte Internationale
du Tapis Vegetal.

The question of scale is of fundamental im-
portance. For the mapping of a given area, the
larger the scale the larger the map and vice-
versa. Alternatively, with a given size of map,
the larger the scale the smaller the area the map
portrays, and vice-versa. A vegetation map of
the largest possible conceivable scale would be
a diagram of a quadrat on which all the individ-
ual plants and their coverage were marked. Such
a map inevitably represents a relatively tiny
area. For the mapping of more extensive areas,
the scale has to be reduced. It then becomes im-
possible to represent individual plants, and they
have to be grouped into communities whose
boundaries are mapped. At progressively smaller
scales fewer communities can be represented and
there has to be progressive generalisation of the
map.

It therefore follows that for the vegetation
mapper there can be no absolute concept of the
plant community. The concept is dictated by
the scale he is using and the ability to represent
units on the map. To give concrete examples,
the vegetation maps of Lord Howe Island
(Pickard 1974) which covers 13 km 2 , of Tas-
mania (Jackson 1965) covering 42 450 km 2 , and
of the Murchison Region of Western Australia
(Beard 1976) about 385 000 km 2 , represent units
of quite a different order from one another, be-
cause of the progressive generalisation which is
obligatory with increasing scale.

It follows also that there can never be any
absolute standard for an inventory of plant
communities of Australia since the number of
communities is dependent upon the scale at
which they are being considered.

Having made a general decision on the use of
the 1 : 1 000 000 scale for principal publication, a
special series of seven map sheets was designed
in order to cover the State in the most econom-
ical manner (Fig. 1). The standard grid of the
International Map of the World Series (also
known in Australia as the Aeronautical Series)
necessitates 14 sheets for Western Australia of
which five straddle the eastern border and are
only half within the State. The special series
however is still built upon the standard grid of
the smaller 1:250 000 series, shown as smaller
rectangles in Figure 1. Basic mapping is done
on aerial photomosaics at a scale of 1:63 360
rather than on individual aerial photographs
because the latter would be too time-consuming.
Adequate detail for the publication scale is
obtained from the mosaics and is first reduced
and drawn at 1:250 000, then reduced again and
compiled at 1:1 000 000. The original 1:250 000
sheets are hand-drawn on film and are filed in
the map library of the Department of Geography
at the University of Western Australia. Dyeline
prints can be obtained on request. Those of the
1:250 000 sheets covering the South-West Bot-
anical Province where vegetation patterns show
finer detail and where there is likely to be greater

public interest, are being drawn on a standard
topographic base and published as a separate
series (Fig. 1.)

Classification and terminology

Having regard to the nature of this survey
which was essentially a preliminary survey of a
very large, little-known area, it was considered
the most appropriate to approach the characteri-
sation of vegetation on a physiognomic basis
(Beard 1973c). Time would not permit the use
of quantitative or phytosociological methods at
this stage, however desirable they might be, and
it was expected that they would be applied in
later, more detailed, work. The broad principles
of classification adopted in this survey, therefore,
are those stated by Beard (1944, 1955). The
basic unit is a floristic unit, the plant association.
The association is the largest possible group with
consistent plant dominants, either of the same
or closely allied species. Associations may be
divided into minor floristic groups and may be
grouped floristically into alliances. Associations
may also be grouped together, according to their
physiognomy (structure and growth-form), into
formations. The formation is thus a physiog-
nomic unit.

Prior to 1964 the physiognomy, classification
and nomenclature of Australian plant formations
had been discussed by Beadle and Costin (1952)
and by Williams (1955) but there was no gene-
rally agreed system. Many advantages seemed
to be offered by the approach suggested by
Kuchler (1949) who drew up a number of simple
life-form classes.

Kuchler’s system had the added advantage
that it could be expressed as a mapping notation.
It is not desirable that a map should depend
entirely on its colours to distinguish units, as
the eye may not readily differentiate between all
of them, and in any case many of the maps have
been published in black and white. The geolo-
gists employ a notation of this sort e.g. Tep
Tertiary, Eocene, Plantagenet Group, Ag— Arch-
aean granite. Vegetation units can be simply
numbered but this is rather a crude approach.

The trouble was that Kuchler’s system, while a
brilliant idea, included illogical features and was
framed to suit North American conditions. Dan-
sereau (1951) reshaped the system into an
elegant and logical classification capable of
describing any vegetation community in the
world. Unfortunately, the mapping notation
became too unwieldy to put onto a map! A
simpler system was therefore devised for practi-
cal use, which has come to be known as the
Beard-Webb system (Beard and Webb 1974). It
was felt that the notation should be of reason-
able brevity and should convey the diagnostic
features of the vegetation which are: 1, Nature
and size of the dominant stratum and of other
strata if of diagnostic importance; 2, Dominant
or diagnostic plant species and 3, Density of
the strata referred to in (1).

77

Published at 1-1 000 000
Published at 1:250 000

In preparation

Figure 1. — Vegetation survey; progress in mapping and publication.

78

Journal of the Royal Society of Western Australia, Vol. 62, Parts 1-4, 1979.

The Beard-Webb system is built up as follows:

(1) Physiognomy of dominant stratum (capital
letters) —

T Tall trees > 30 m tall.

M Medium trees — 10-30 m tall.

L Low trees < 10 m tall.

S Shrubs > 1 m tall.

Z Dwarf shrubs < 1 m tall.

G Bunch grasses.

H Hummock grass (spinifex).

F Forbs.

X Lichens and mosses.

C Succulents.

(2) Floristic (lower case letters) —
e Eucalyptus.

a Acacia,
t Triodia.

x Heterogeneous (mixed or other).

(3) Density (lower case letters) —

d Dense canopy. Projective foliage cover,
> 70 per cent.

c Mid-dense canopy. Projective foliage cover,
30-70 per cent.

i Incomplete canopy, open not touching.

Projective foliage cover, 10-30 per cent,
r Rare but conspicuous. Projective foliage
cover, < 10 per cent.

b Barren, vegetation largely absent. Pro-
jective foliage cover negligible,
p Scattered groups. No definite foliage cover.

The letter “C” has been introduced to the
classification and defined as “succulents”. Most
of these are Chenopodiaceae, hence the letter
“C”, “S” being pre-occupied. In addition to the
fully succulent samphires, this class is intended
to include the semi-succulent saltbush and blue-
bush vegetation, which is locally very widespread
in parts of the Australian Eremaea, as it is con-
sidered essential that these communities be
recognized to have a unique character of their
own, and differentiated from sclerophyll shrub -
lands. The floristic component (e, a, t, etc.)
can be differentiated as ei, e 2 to indicate par-
ticular species, and other letters may be added
as required, e.g. m. for Melaleuca.

The actual formulae are to be written with the
floristic category first e.g. eMc, aLi: meaning
respectively a eucalypt-dominated closed medium
tree community (a eucalypt forest in fact) and
an acacia-dominated woodland (mulga). The
formulae are designed in this “triangular” form
in order to be more readily comprehended at a
glance. The central capital letter conveys the
most important feature (physiognomy), the left-
hand one floristics and the right-hand one den-
sity.

The principal, capital-lettered category is
based upon Kuchler’s Group 1 Height, and is
intended to accord mention primarily to the
dominant stratum, e.g. eMc. If two or more
strata are considered co-dominant, their symbols
may be written together e.g. xSZc. Any diagnos-
tically important strata may be included in this
manner — eLr, aSr, pHi.

The floristic category serves to name the
species or genera which are dominant or diagnos-
tic, but as most West Australian communities

are of simple composition or even single-dom-
inant associations, this category will also to a
large extent convey — from the known morph-
ology of the species — a growth -form character-
istic of the community. A category “hetero-
geneous” is provided for mixed communities in
which no definite dominance asserts itself.

The third category, density, is the same as
Kuchler’s Group II. KUchler’s Group III is not
directly included: the characters which it deals
with will have been already incorporated in the
formula if important i.e. if they are “salient
features” otherwise they are disregarded. Dan-
sereau has set up three other categories in his
system: function, leaf shape and size, and leaf
texture. All of these will be found to be con-
veyed by the floristic category and are therefore
not separately required.

Specht’s projective-foliage-cover classes have
been introduced and related to the code letters,
with the addition of a letter “d” for “Dense” not
appearing in Kuchler or in Beard (1969). It is
intended in Australian work that “d” should
apply to rain forests while “c” will be used for
eucalypt forests which admit much more light
even though tree crowns may be touching.

An attempt is being made to carry the treat-
ment in the notation and terminology through
into the map colour scheme. A colour spectrum
ranging from blue for moist forest through to
red for desert is in accord with general inter-
national practice and has been planned as fol-
lows:

T Blue
M Green
L Orange
S Yellow
Z Grey
G Olive
H Red

F Rare, not provided
X Hachured
C Brown

Density of the vegetation is conveyed by shade
of colour, dark for dense, pale for open. The
general transition from the desert southwards is
expressed in the sequence of colours red-orange-
yellow-green-blue, and northwards in red-yellow-
olive-green. “Z” and “C” are mainly edaphi-
cally controlled and are therefore extraneous to
the climatic sequence. Considerations of map
design are involved at this stage and the colour
sequences are designed to be aesthetically pleas-
ing.

Table 1 shows the Beard-Webb classification
with its associated nomenclature. It differs from
the rather similar treatment of Specht (1970) in
basing the classification upon the dominant
stratum, not the tallest one, and is more in
accord with ecological principles as well as being
more suited to mapping. The second part of the
table provides for wooded grass and shrub com-
munities which have more than one significant
layer. Grassland nomenclature follows Beard
(1966b).

79

Journal of the Royal Society of Western Australia, Vol. 62, Parts 1-4, 1979.

Table 1

Nomenclature of vegetation units as used in the vegetation survey of Western Australia
A. Communities with a single significant layer

Cover of dominant stratum

Life Form/Height Class

Dense d

70-100%

Mid-dense c

30-70%

Incomplete i

10-30%

Sparse r

<10%

Very Sparse b

Negligible

T Tall trees >30 m

Dense Tall Forest

Tall Forest

Tall Woodland

Open Tall Woodland

M Medium trees 10-30 m

Dense Forest

Forest

Woodland

Open Woodland . ..

L Low trees < 10 m ....

Dense Low Forest

Low Forest

Low Woodland

Open Low Woodland

Sparse Low Wood-
land

S Shrubs > 1 m

Dense Thicket ....

Thicket

Scrub

Open Scrub

Sparse Scrub

Z Dwarf shrubs < 1 m

Dense Heath

Heath

Dwarf Scrub

Open Dwarf Scrub

Sparse Dwarf Scrub

G Bunch Grasses

Dense Grassland

Mid-dense Grassland

Grassland ....

Open Grassland ....

Sparse Grassland

H Hummock grasses ....

Hummock grassland

Open Hummock
grassland

Sparse Hummock
Grassland

F Forbs

Dense Herbfield

Mid-dense Herbfield

Herbfield

Open Herbfield

Sparse Herbfield

X Lichens and mosses

Mat Plants

Open Mat Plants ....

Sparse Mat Plants

C Succulents .. .

Succulent Steppe

Open Succulent
Steppe

Sparse Succulent
Steppe

B. Communities with more than one significant layer

Cover of tree/shrub layer

Description

Incomplete i

10-30%

Sparse r

<10%

Very Sparse b

Negligible

Absent

Wooded Bunch Grassland

Savanna Woodland ....

Tree Savanna
Shrub Savanna

Sparse Tree Savanna
Sparse Shrub Savanna

Grass Savanna

Wooded Hummock Grassland

Steppe Woodland

Tree Steppe

Shrub Steppe

Sparse Tree Steppe ....
Sparse Shrub Steppe

Grass Steppe

Wooded Succulent Steppe ....

Thickly Wooded
Succulent Steppe

Lightly Wooded
Succulent Steppe

Sparsely Wooded
Succulent Steppe

Succulent Steppe

Heath with Trees

Tree heath

Heath with shrubs

Scrub heath

Heath with mallee

Mallee health

Progress of publication by the Vegetation Sur-
vey of Western Australia up to the time of writ-
ing (August 1978) is summarised in Figure 1. Five
of the seven 1:1 000 000 map sheets have been
published (Beard 1974, 1975b, c, 1976a, Beard and
Webb 1974) and all but eight of the 1:250 000
map sheets are available (Beard 1969, 1972 a-e,
1973 a, b, 1976 b-f, Smith 1972, 1973, 1974). The
survey is thus almost complete and has lifted
Western Australia from the position of least to
best documented of the States in terms of vege-
tation.

Future work

It is considered a desirable objective that cov-
erage of Australia at the 1 : 1 000 000 scale should
be completed as soon as possible. Tasmania was
done in 1965 (Jackson 1965). State vegetation
mapping projects have been commenced at the
National Herbarium, Sydney, and at the Queens-
land Herbarium, but results have not yet been
published. It is also a desirable objective that
the mapping should be uniform. The authorities

conducting geological surveys in Australia long
ago decided to adopt uniform mapping standards
for the general 1:250 000 series so that maps of
any State are comparable. The questions of
scale come up again here, however. The stand-
ards of the vegetation survey of Western Aus-
tralia, the Beard-Webb classification system, the
related map colours and so on, apply to the
1:1 000 000 and 1:250 000 scales for which they
were devised. Uniformity at other scales is not
suggested at this stage. Carnahan’s (1976) map
of Australia at 1:6 000 000, used a modified sys-
tem based upon Beard-Webb, but changed to
suit the requirements of his scale. More inten-
sive, larger-scale, work would also have to set
its own standards. This can already be seen
happening in the work of B. G. Muir (1977)
in the Western Australian Museum’s “Biological
Survey of the W.A. Wheatbelt”, where
the Beard-Webb system was expanded by the
introduction of several new Life-Form-Height
classes to provide for a more intensive definition
of stratification. Thus Low Trees have been
divided into two, LA 5 to 15 m tall, and LB

80

Journal of the Royal Society of Western Australia, Vol. 62, Parts 1-4, 1979.

< 5 m; a class for the mallee life-form has been
introduced and given the letter K, divided into
both tree and shrub forms; shrubs are divided
into five height classes, bunch grasses into two,
while additional classes have been introduced for
mat plants (M), herbaceous species (J), sedges
(V) and cryptogams (X).

This development is regarded as both logical
and welcome: it was always envisaged that some-
thing of this kind should and would take place
when others came to hang more intensive work
within the framework of the extensive survey.
It serves to emphasise the point that scale*
dictates procedure.

Some other large-scale mapping has been
carried out recently in Western Australia mainly
for biological surveys and should be mentioned,
e.g. Beard 1967, 1976g, McKenzie, Burbidge and
Marchant 1973, Dames and Moore, 1975, Smith
1977. A small-scale map of the State at
1:10 000 000 has been prepared by reduction of
the 1:1 000 000 sheets, recognising 25 basic vege-
tation units (Beard 1979a).

Derived benefits of the vegetation survey

Vegetation mapping was used for a study of
palaeodrainage in the arid zone of the State, in
which a map showing the approximate courses
and directions of rivers which must have existed
in more pluvial times was produced, Beard
1973d). The general accuracy of the interpreta-
tion has been confirmed by later work (van de
Graaff et al. 1977).

Vegetation mapping has also been used to
define phytogeographic natural regions in the
State. Diels (1906) in treating Western Austra-
lia south of the tropic divided the area into two
Botanical Provinces, the South-West Province
and the Eremaean Province, and subdivided the
former into 6 Botanical Districts, the latter (as
far as known to him) into 2. Gardner (1942)
added the concept of a Northern Botanical Pro-
vince in the tropics. He later expanded Diel’s
Botanical Districts on a State-wide basis
(Gardner and Bennetts 1956) so that there were
now 5 districts in the Northern Province, 5 in the
Eremaean, and 6 in the South-West. Burbidge
(1960) incorporated the Botanical Provinces into
Australia-wide Floristic Zones, and treated the
Coolgardie Botanical District as an Interzone or
ecotonal region lying between the Eremaean and
South-West Provinces.

When vegetation mapping commenced it
quickly became apparent that natural regional-
isations were present in the grouping of vegeta-
tion units and were expressed by the map colours.
To a very large degree these coincided with the
intuitive ideas of Diels and Gardner and the map-
ping was therefore used as a means of laying
down precise boundaries for the Botanical
Provinces and Districts previously rather arbit-
rarily sketched on a small scale. The regional
boundaries have been marked on the vegetation
maps, and a new State-wide small-scale map has
recently been produced (Beard 1979b).

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North Kimberley, W.A., eds. E. D. Kabay and
A. A. Burbidge, Wildlife Research Bull 6.
Smith, R. (1951). — Soils of the Margaret River-Lower
Blackwood River Districts, Western Austra-
lia. CSIRO Aust. Bull. 262.

Specht, R. L. (1970). — Vegetation. In The Australian
Environment, ed. G. W. Leeper. CSIRO and
MUP, Melbourne.

Speck, N. H., Bradley, J., Lazarides, M., Patterson, R. A.,
Slatyer, R. O., Stewart, G. A. and Twidale,

C. R. (1960). — Lands and Pastoral Resources
of the North Kimberley area, W.A. CSIRO
Aust. Land Res. Ser. 4.

Speck, N. H., Wright, R. L., Rutherford, G. K., Fitzgerald,
K., Thomas, F., Arnold, J. M., Basinski, J, J..
Fitzpatrick, E. A., Lazarides, M. and Perry
R. A. (1964). — General Report on Lands of
the West Kimberley area, W.A. CSIRO Aust.
Land Res. Ser. 9.

Stewart, G. A., Perry. R. A., Paterson, S. J., Traves,

D. M., Slatyer, R. O., Dunn, P. R., Jones,
P. J. and Sleeman, J. R. (1970) — Lands of the
Ord-Victoria Area W.A. and N.T. CSIRO Aust.
Land Res. Ser. 28.

Van de Graaff, W. J. E., Crowe, R. W. A., Bunting, J. A.,
and Jackson, M. J. (1977) — Relict early
Cainozoic drainages in arid Western Austra-
lia Z. Geomorph. N.F. 21: 379-400.

Williams, R. F. (1955). — Vegetation Regions. Atlas of
Aust. Resources, Dept. Nat. Development,
Canbella.

Wood, J. G. (1937). — The Vegetation of South Australia,
Govt. Printer, Adelaide.

82

Journal of the Royal Society of Western Australia, Vol. 62, Parts 1-4, 1979, p. 83-94.

Aspects of Science in Western Australia 1829-1979.

A volume to commemorate the 150th Anniversary of the founding of Western Australia.

Plant physiologist amongst the grain legumes

by J. S'. Pate

Botany Department, University of Western Australia, Nedlands, W.A. 6009

Abstract

This paper expresses a physiological approach to plant functioning based on
studies of the efficiency of operation of plant organs and whole plants. Applied
to nodulated legumes, it has the purpose of discovering how efficiently photosynthetic
resources are utilized for nitrogen fixation and the synthesis of protein in vegetative
parts and seeds. Comparisons between species suggest how differences in structure
of organs and in growth habit and reproductive behaviour of the legume may
influence its efficiency of functioning and yield of seed. The techniques illustrated for
studying the economy of carbon and nitrogen might well be applied widely in studies
of the productivity of non-leguminous crops.

Introduction

According to figures published by the Food
and Agricultural Organization of the United
Nations (FAO) the annual world yield of cereals
and grain legumes amounts to approximately
1 500 million tonnes of harvested seed, the
equivalent of three-quarters of the food dry
matter used for human consumption. Although
production of cereals within this total is more
than ten times greater than that of legumes, the
high protein content of legume seeds (Table 1),
and the high levels of certain essential amino
acids in their seed protein make legume grain
uniquely important in the nutrition of man and
his animals, especially when complementing a
predominantly cereal diet (Siegel and Fawcett
1976).

Despite their importance to man most species
of grain legumes have been subjected to much
less extensive research than have cereals and
other major crop plants. This becomes apparent
when examining the volume of literature
generated annually on different crops, the extent
of national and international funding for
research on leguminous versus non-leguminous
crops, and the deployment of scientific personnel
within research institutes concerned with the
breeding and physiology of agriculturally-
important plants.

The sad state of grain legumes on the world
scene is also obvious when one follows changes
in their productivity over the past twenty years,
as indicated by yield figures given by the FAO.
During this period average yields of cereal crops
per hectare have risen by some 65%, due
especially to the development of high-yielding
cultivars of rice, wheat and maize, but little
or no improvement is evident over the same
period in productivity of most of the major
grain legumes. Even in those species subjected

to intensive selection programmes, for example
soybean (Glycine max (L) Merr.) and garden
pea (Pisum sativum L.) the improvement in
yield per hectare over the 20-year period has
been modest — of the order of 20-35%. This
situation, combined with the low baseline
productivity of legume crops as a whole (Table
1), the uncertainty of yield realization from one
season or soil type to the next, and the notorious
susceptibility of grain legumes to diseases, pests
and environmental stress, would make one doubt
whether further increases in funds and
manpower should be devoted to their exploita-
tion as crop plants.

Despite this distressing situation much can
still be marshalled in defence of grain legumes.
Firstly, it may be argued that their usefulness
and productiveness should be assessed primarily
in terms of protein yield. When viewed on
this basis the comparisons with cereal crops
become much more favourable, with several
legume species outyielding cereals in protein
per hectare by a considerable margin (Table 1).
Moreover, in the case of peanut (Arachis
hypogaea L.) and soybean a second valuable
product, oil, is present in large amount in the
seed. Secondly, the role of legumes in crop
rotations should be recognized. In many parts of
Asia, for example, grain legume crops are often
grown at times of the year not suitable for
cereals and under climatic conditions inferior to
those allotted to the cereal. Lower yields from
the legume are to be expected under these
conditions, but, the fact that a second,
nutritionally-valuable crop can be included in
the year’s cultivation, commends use of the
legume. Thirdly, many grain legumes are short-
cycle crops, occupying the ground for a much
shorter period of time than in the case of
many cereal crops; again comparisons based on
yield of seed per hectare are unfair. Finally,

83

Journal of the Royal Society of Western Australia, Vol. 62, Parts 1-4, 1979.

Table 1

World production and average yields of major grain legumes and cereals

Grain legumes

Pro-

duction

1974

(million

tonnes/yr)

Average

yield

1965-74

kg/ha

Protein
level in
dry weight
of seed

%

Average

seed

protein

yield

1965-74

kg/ha

Major producing countries (in order
of importance as producers) 1974*

Soybean ( Glycine max (L.)Merr.)

56-8

1 295

38-5

499

USA, China, Brazil, Indonesia, Mexico,
USSR, Canada

Peanut (in shell) ( Arachis hypogaea L.)

17-6

879

28-6

251

India, China, Nigeria, Senegal, Sudan,
Burma, USA

Dry beans ( Phaseolus spp.)

115

484

260

126

Brazil, India, China, Africa, USA, Mexico,
Europe

Dry pea ( Pisum sativum L.)

11-7

1 117

230

257

USSR ,t China, W. Europe, USA, Ethiopia,
S. America

Chickpea ( Cicer arietinum L.)

5-9

617

19 9

123

India, Pakistan, Mexico, Ethiopia, Turkey,
Burma

Broad bean ( Vicia faba L.)

5-2

1 076

27-5

296

China , Italy, Egypt, U.K., Morocco, Ethi-
opia, Brazil

Vetches ( Vicia spp. Lathyrus spp.)

21

1 241

27-5

341

USSR. Turkey, Spain, Poland, Morocco,
Greece

Pigeon pea ( Cajanus cajan (L.) Mi lisp.)

1-5

653

22-1

144

India, Africa, Latin America

Cowpea ( Vigna unguiculata (L.) Walp.)

11

330

26-4

87

Nigeria, Upper Volta, Uganda, Asia, USA

Lentil ( Lens esculenta Moench.)

1-2

609

24-5

149

India, Turkey, Ethiopia, Syria, USSR, W.
Europe

Lupin ( Lupinus spp.)

0-8

828

34-5

286

USSR, Poland, Australia, W. Europe

Others

3-3

522

Total legumes

118 7

Total cereals t

1-357

1 750

11

193

* Calculated from average values for seed protein content given in the Table.

t Where the name of a country is printed in italics its contribution accounts for more than half of the world’s production.
t Includes wheat, barley, maize, sorghum and rice.

the unique contribution made by legume crops
to soil fertility through their nitrogen-fixing
symbiosis with Rhizobium , must be recognized.
According to data provided by Nutman (1976)
this can amount to 60-210 kg N per hectare
per growing season. The attendant savings in
reduced input of synthetic nitrogen fertilizers
are likely to be quite substantial, whether in
terms of expense to the farmer, or, indirectly,
as a reduction in run-off of nitrogenous
fertilizer residues into surrounding ecosystems.

The challenge then to students of the grain
legumes is to assess from a variety of viewpoints
the future value of these plants on the world
agricultural scene. Agronomists will contribute
by indicating how better management practices
can lead to higher seed yield under a range of
soil types and environments. Plant breeders
will contribute by developing high-yielding
cultivars resistant to disease, adapted to specific
environments, and productive of better quality
protein. The research of plant physiologists is
likely to provide a fundamental understanding of
how legumes function and an indication of the
processes likely to limit production of seed
protein under a range of agricultural conditions.

Questions relating to the productivity of
plants require detailed knowledge of the
integrated functioning of the whole plant and
its parts and investigations of the vast spectrum
of related events and processes at structural,
physiological and biochemical levels which
accompany development of the plant and its
organs during growth. The purpose of this
paper is to highlight this type of approach as
applied to grain legumes, concentrating
particularly on studies conducted by the author
and his colleagues in Belfast, Northern Ireland
and currently at the University of Western
Australia. The first section of the paper deals
with investigations of the functioning of
nodulated roots, especially in relation to their
utilization of carbohydrate for nitrogen fixation.
The second section concerns the developing
legume fruit and the mechanisms involved in its
conversion of imported assimilates into the
reserves of seeds. The third section is an
examination of the functioning of the whole
grain legume, in an attempt to visualize how it
deploys its photosynthetic product for vegetative
growth, nitrogen-fixation, and the storage of
protein in seeds.

84

Journal of the Royal Society of Western Australia, Vol. 62, Parts 1-4, 1979.

The functioning of nodulated roots

The primary function of plant roots is to
supply minerals, water, and certain synthetic
products to the shoot system. In legumes the
synthetic functions of below-ground parts
assume unusual importance, due to the nitrogen-
fixing activities of nodules or the assimilation
in roots of combined forms of nitrogen such as
ammonium or nitrate (Pate and Wallace 1964;
Wallace and Pate 1965; Oghoghorie and Pate
1971).

It is now generally agreed that ammonia is
the first stable product formed during nitrogen
fixation by the bacteroids of the nodule
(Bergersen 1971; Dilworth 1974), and that its
assimilation into organic compounds probably
takes place via the glutamate synthase/gluta-
mine synthetase system of the host cytoplasm
of the bacterial tissue (Miflin and Lea 1977).
From the glutamine produced by this system,
other secondary products form, these and gluta-
mine being exported from the nodule to the plant
shoot through the transpiration stream (Pate
1971).

Decapitated roots and detached nodules of
grain legumes commonly bleed a fluid from their
xylem, and this can be readily collected and
analysed to reveal which compounds are being
utilized for export of fixed nitrogen. Some
grain legumes (e.g. Lupinus spp., Pisum spp. and
Vicia spp.) export upwards of 70% of their fixed
nitrogen as the amides glutamine and aspara-
gine, the latter compound usually predominating
(Pate et al. 1969; Gunning et al. 1974), whilst
in other legumes (e.g. Phaseolus spp., Vigna
unguiculata (L.) Walp.) the ureides allantoin
and allantoic acid are the principal exports
(Atkins et al. 1978). These various nitrogen-
ous products are readily labelled with 14 C and
51 N following feeding of the legume shoot with
14 C0 2 and its root gas space with 15 N 2 (Pate 1962;
Oghoghorie and Pate 1972; Herridge et al. 1978),
indicating that the compounds in question are
true metabolic products of the fixation process.

Roots are of course dependent on carbohy-
drates received from the shoot as translocated
sugar. Some of this carbohydrate is consumed
in respiration of the nodulated root, some is used
in formation of dry matter during growth of
roots and nodules, and some serves as carbon
source for formation of the fixation products
which ultimately leave the root in the transpira-
tion stream. The extent of these different forms
of usage of carbonhydrate has been assessed
during growth of two legumes, white lupin ( hup -
inus albus L.) (Pate and Herridge 1978) and
cowpea (Vigna unguiculata) (Herridge and Pate
1977). Carbon incorporated during growth is
assessed by measurements of dry weight gains
of root and nodules over specific intervals of
growth. The respiratory output of the enclosed
nodulated root of the intact plant is measured
continuously by a gas-flow system involving
collection of respired C0 2 in Pettenkoffer assem-
blies (Minchin and Pate 1973), and the propor-
tion of this C0 2 efflux due to nodules is estimated
by comparing the specific activity of respiration

of freshly detached nodules with that of their
supporting root. Finally, the types and concen-
trations of compounds in xylem exudate of the
nodulated root are assessed, and, by relating the
C:N ratio of these exports to the total nitrogen
increment of the shoot, the amount of carbon
cycled through the root as fixation products is
estimated.

In both cowpea and white lupin the carbon
used in export of fixed nitrogen comprises the
largest item of the carbon budget of the nodule,
respiratory losses from the nodule the next
largest item, and the incorporation of carbon
into nodule dry matter the least expensive item.
The carbon balance of the whole nodulated roots
shows root respiration to be the largest consumer
of carbohydrate. The large tap root of lupin
is clearly more expensive to maintain than the
less extensive root system of the cowpea. When
their consumption is compared in relation to
how much nitrogen they fix, nodules of lupin
are found to use carbon less efficiently than those
of cowpea (Table 2; Fig. 1). One reason for this
is the higher C:N ratio of the amides exported
from lupin nodules than of the ureides exported
from cowpea nodules, but C0 2 efflux per unit
of nitrogen fixed is also greater from nodules of
lupin than of cowpea (Table 2), suggesting that
other elements are involved in efficiency of
operation.

Table 2.

Economy of carbon in nitrogen- fixing nodules of
two grain legumes*

Vigna

Lupinus

Species

unguiculata albus

days

Period of growth

0.78t

0.94f

mg. plant- 1

N fixed

726

788

N exported

.... 705

761

C used in export of N. ±

969

1 789

C lost as C0 2 in nodule respiration 789

1 372

C incorporated into nodule mass

.... 253

298

Total C requirement of nodule

.... 2 011

3 459

g atom

. g atom- 1

Total C required/N fixed

.... 3.23

5.12

C exported/N exported

.... 1.60

2.74

C respired/N fixed

.... 1.27

2.02

* Data from Herridge and Pate 1977; Pate and Herridge
1977; Atkins, Herridge and Pate 1978.
f Vegetative growth and early fruiting.
t Estimated from the amount of N exported and the
C:N ratio of xylem sap.

Since nodules have a relatively high rate of
consumption of translocate in proportion to their
mass, their functioning in nitrogen fixation is
especially vulnerable to a decline or interruption
in supply of photosynthate from the shoot. How-

85

Journal of the Royal Society of Western Australia, Vol. 62, Parts 1-4, 1979.

ever, they appear to possess a strategy of diurnal
functioning in which sugar and polysaccharide
built up during the day serve as carbon sources
for fixation at night, thus allowing the nodule
to fix much more nitrogen at night than would
be possible were it to have to rely solely on cur-
rent translocate (Minchin and Pate 1974). In
garden pea ( Pisum sativum L.) and cowpea,
removal of the shoot causes within 24-48 hours
a marked decline in the nitrogen-fixing activity
of the root, as gauged by acetylene-reducing
performance (Halliday 1976), but in broad bean
(Vicia faba L.) or white lupin, fixation continues
unabated for several days after removal of the
shoot, suggesting that carbohydrate reserves of
the tap-root might be made available as a sub-
strate for nitrogen fixation. Drought-induced
defoliation can also cause root reserves to be
utilized for nitrogen fixation (unpublished data) .

It has recently proved possible to compare
experimentally-obtained values for C0 2 efflux of
nodules with theoretical estimates of the costs,
in terms of respired C0 2 , of providing the ATP
and reductant necessary for converting N 2 to
NH 3 , and for assimilating this ammonia into" the
compounds known to be exported from the
nodule (Atkins et al. 1978). A summary of
the relevant calculations (Table 3) shows that,
for the three grain legumes studied, estimates of
C0 2 output based on the theoretical reauirements
for ATP by the nitrogenase system and ammonia
assimilating reactions exceed by a substantial
margin the experimentally-observed C0 2 output

of the nodules. The differences involved appear
to be greater in one legume (cowpea) than in the
other two (garden pea and white lupin). The
discrepancy between observed and predicted C0 2
efflux would be partly resolved were the nodule
to economise in its carbon usage by refixing some
of its respired C0 2 , and to utilize this carbon in
synthesis of the carbon skeletons of the com-
pounds employed in transporting fixed N to the
shoot. The demonstration of an active phos-
phoenol-pyruvate carboxylase system in nodules
of Lupinus angustifolius L. by Christeller et al.
(1977) suggests that nodules assimilate C0 2 , and
do so at a sufficiently fast rate (up to 0.8 mole-
cules of C0 2 incorporated per atom of N fixed)
to effect a considerable saving in terms of im-
ported carbon from the shoot. It remains to be
seen whether this applies to legumes generally.

The most likely source of discrepancy between
observed and predicted values for C0 2 output
from N-fixing nodules relates to the uncertainty
of the in vivo requirements of the nitrogenase
for ATP. The estimates of Table 3 assume that
4 ATP molecules are needed per electron pair
utilized by nitrogenase, a value regarded as an
average estimate for the performance of isolated
nitrogenase in vitro (Dil worth 1974). In living
bacterial tissue nitrogenase might operate at
much lower cost since the structural configura-
tion of the enzyme system in vivo might not
require the continuous stabilization involving
ATP hydrolysis needed to maintain activity of
the extracted enzyme (Atkins et al. 1978).

86

Journal of the Royal Society of Western Australia, Vol. 62, Parts 1-4, 1979.

Table 3.

Theoretical estimates of and experimental observations
on CO 2 production by nitrogen- fixing nodules of
three grain legumes.

Vigna

Lupinus Pisum

(A)

Theoretical Estimates <0

mol

ATP . mol NH 3- 1

1 .

ATP requirements for syn-
thesis of organic solutes of

nitrogen from NH 3 ( 2 ) —
amino acids

0.40

0.84

0.48

glutamine-amino

0.20

0.24

0.30

glutamine-amido

0.05

0.06

0.08

asparagine-amino

0.20

1.34

1.26

asparagine-amido

0.15

1.01

0.95

ureide

Total cost for NH ;i assimila-

2.10

—

0.30

tion

3.10

3.49

3.37

2 .

C0 2 output associated with

mol C0 2 . mol

NH 3-1

nitrogen assimilation < 3 >

N 2 reduction to NH ;J

NH 3 incorporation into

1.75

1.75

1.75

organic N solutes

0.52

0.58

0.56

Total N assimilation cost ....

2.27

2.33

2.31

3.

C0 2 output for growth and

mol

C0 2 . mol NH 3- 1

maintenance.

C loss as C0 2 in nodule

formation (■*)

C loss as CO -2 in nodule

0.06

0.06

0.11

maintenance

0.05

0.06

0.01

Total nodule cost

0.11

0.12

0.12

4.

Total C0 2 output.

mol

C0 2 . mol

NH 3- 1

Items 2 and 3 ....

2.38

2.45

2.43

(B) Experimental Observations.

Output of respired C0 2 from

mol

C0 2 . mol NH 3- 1

nodule

1.27

2.02

1.73

(1) As detailed by Atkins et al. (1978).

(2) Synthetic pathways as described by Atkins et al.
(1978), proportions of amino compounds produced
as suggested from xylem sap analysis.

(3) Calculated as 6 ATP per CO 2 , assuming that the
P/2e- ratio of oxidative phosphorylation is 3.

(4) Calculated using 0.2 g C0 2 produced . g dry wt- 1
synthesized (Penning de vries et al. 1974).

(5) Calculated using 30 mg glucose . g dry matter -1
day- 1 (from Penning de Vries 1975).

(6) From studies of Minchin and Pate 1973 (Pisum)-
Herridge and Pate 1977 ( Vigna ); Pate and Herridge
1978 (Lupinus).

An Important factor in the efficiency of func-
tioning of nitrogenase relates to its activity in
converting protons to hydrogen gas, a process
which will compete for electrons with the reduc-
tion of nitrogen gas to ammonia (Dixon 1975).
In nodules formed by certain legum e-Rhizobium
associations hydrogen evolution is high relative
to ammonia formation; in others a hydrogenase
system re-utilizes much of the hydrogen which
is produced, the attendant conversion to water
generating significant amounts of ATP (Dixon
1975; Schubert and Evans 1976). It follows that
nodules possessing an active hydrogenase system
will have a lower requirement for ATP than
those not so endowed, and that their overall
consumption of carbon in nitrogen fixation will

be correspondingly reduced. It appears that the
strain of Rhizobium, not the host plant, is
responsible for determining the level of hydro-
genase in the nodules of a particular association
(Schubert et al. 1977) but that ontogenetic and
environmental factors may modify H 2 evolution
from nodules during the growth cycle of a speci-
fic association (Bethlenfalvay and Phillips 1977).
This suggests that nodules may not operate at a
constant level of hydrogenase activity relative to
nitrogen fixation. There is still considerable con-
fusion surrounding the significance of hydrogen
evolution and hydrogenase activity in nodule
functioning, but it would be especially interesting
to see whether the apparent differences in effici-
encies of carbon usage by the N-fixing associa-
tions in Table 2 relate to effects of this nature.

The structure and functioning of fruits

Grain legumes display great variation in size,
shape and structure of their fruits. In some
(e.g. peanut ( Arachis hypogaea) and bambarra
ground nut ( Voandzeia subterranea Thon.)) the
fruits develop underground and lack chlorophyll.
In most, however, the fruits are green and are
borne aerially, often in a well-illuminated situa-
tion in the canopy. They thus have the poten-
tial for photosynthesis and for attracting water
and nutrients from the parent plant through
their transpirational activity.

The legume fruit consists of a single carpel
containing 1-10 or more seeds. Its vascular
network comprises a ventral longitudinal strand,
two dorsal longitudinal strands supplying the
seeds, and a network of veins traversing the
lateral walls of the pod and connecting above
and below with the longitudinal strands. A gas
cavity of variable size and shape forms within
the carpel as the young fruits develop. During
later growth this gas space is encroached upon
by the developing seeds (Harvey et al. 1976;
Flinn et al. 1977).

Anatomists recognise three basic layers in the
pod wall of legumes — exocarp, mesocarp and
endocarp (see Esau 1965; Fahn 1974; Fig. 2).
The exocarp comprises a heavily-cutinized epi-
dermis, poorly supplied with stomata, and, in
some genera, an underlying hypodermis of ligni-
fied cells. The mesocarp consists of 5-40 layers
of thin-walled, highly vacuolated cells, usually
well supplied with plastids. The outer mesocarp
contains the bulk of the chlorophyll and displays
prominent air-spaces, especially adjacent to
stomata (Fig. 2). The inner mesocarp shows
few, if any, air spaces and contains the vascula-
ture of the pod wall. The endocarp consists of
one or more layers of sclerenchyma, an inner
epidermis, and, in some genera, an intervening
zone of parenchyma. Since cultivated legumes
have been selected for non-shattering pods the
fibrous component of the endocarp is usually
much reduced in comparison with ancestral
forms of the species (Zimmermann 1936). In
certain grain legumes, for example the “mange
tout” forms of garden pea, sclerenchyma may be
completely absent, making the fruits soft and
edible.

87

Figure 2. — Pod wall structure in grain legumes. (A) Transverse section of the pod of Pisum sativum L. showing
wall layers (xl95). (B) Endocarp of pod of Lupinus albus L. (x430). (C) Endocarp of pod of Pisum sativum

(x850). (D) Electron micrograph of portion of cell of photosynthetic inner epidermis of the pod of Pisum

sativum showing chloroplast (x24 000). OE — outer epidermis; SG — starch grain; VB — vascular bundle; V —
inward projecting veinlet of vascular network; F — fibre layer; P — parenchyma of endocarp; IE — inner epi-
dermis (photosynthetic); CP — chloroplasts of inner epidermis; M — mitochondria; MB — microbody; CW — cell
wall. (Photographs kindly supplied by Dr. J. Kuo, Botany Department, University of Western Australia).

88

Journal of the Royal Society of Western Australia, Vol. 62, Parts 1-4, 1979.

The pod walls of grain legumes are far from
opaque, with up to half of the photosyntheti-
cally-active radiation received at the pod sur-
face being transmitted to the inner epidermis
(Atkins et al. 1977). In many genera (e.g.
Lupinus, Pisum, Vigna , Glycine, Cajanus, Cicer )
the cells of the inner epidermis are equipped with
chloroplasts (Kuo and Pate unpublished; Fig. 2)
and in some cases (e.g. Pisum sativum and
Lupinus albus ) the free-ending veinlets of the
vasculature of the pod wall project inwards
from the main vasculature to terminate almost
at the surface of the endocarp, an adaptation,
one might suppose, for collection of assimilates
produced by the photosynthetic endocarp. In
Lupinus albus these veinlets consist solely of
phloem (Kuo and Pate unpublished) a feature
consistent with a retrieval function within the
pod wall.

Several physiological studies on grain legume
fruits have provided information directly rele-
vant to pod structure. Rates of exchange of C0 2
and transpiration rates are low, in comparison
with leaves, as suggested by the structural
features of the outer epidermis of the pod (Flinn
and Pate 1970; Crookston et al. 1974; Andrews
and Svec 1975). Poor ventilation also results
in high concentrations of C0 2 (0.05-2.5% by
volume of the fruit’s gas space) (Harvey et al.
1976; Atkins and Pate 1977), much of this C0 2
being attributed to respiration of the developing
seeds. Since C0 2 levels in the gas space are
lower in illuminated than in darkened fruits,
photosynthetic recycling of carbon by the inter-
nal tissues of the fruit is suspected. In Pisum
sativum, a species with chloroplasts in its inner
epidermis, injection of 14 C0 2 into the gas cavity
of an illuminated fruit leads to labelling of the
pod but not the seeds, but neither pod nor seeds
become labelled if the 14 C0 2 injected into a fruit
in darkness (Flinn et al. 1977). Labelled assimi-
lates formed from 14 C0 2 injected into an illumin-
ated fruit can be traced first to the vascular
network of the pod and then to the seeds, con-
firming the pod’s role in recycling respired car-
bon back to the seeds.

Up to 25% of the chlorophyll of the pod of
garden pea is contained in the inner epidermis,
and this single layer also contains significant
fractions of the pod’s C0 2 -assimilating enzymes,
ribulose diphosphate carboxylase and phosphoen-
olpyruvate carboxylase (Atkins et al 1977). The
last of these two enzymes may function par-
ticularly well in the C0 2 -rich environment of the
endocarp, and it is significant that the same
enzyme occurs in high concentration in seeds of
certain grain legumes (Hedley et al. 1975; Atkins
and Flinn unpublished data).

Using data for compositional changes in pod
and seed during growth, information on the day
by day exchange of C0 2 and transpiration of the
fruit, and analysis of the xylem and phloem
fluids supplying the fruit with water and assimi-
lates, it has proved possible to construct a quan-
titative model of a legume fruit’s functioning
based cn its intake of carbon, nitrogen and water
through xylem and plPoem. The model for
white lupin, as described by Pate et al. (1977)
(see Fig. 3), indicates that 93% of the imported

xylem

Figure 3. — Economy of carbon, nitrogen and water in a
fruit of lupin (Lupinus alba L. cv Neutra). Deliv-
eries through xylem and phloem are expressed rela-
tive to a net intake of 100 units of a specific com-
modity. Net photosynthesis, respiration loss, trans-
piration, water losses from tissues during fruit
ripening and nitrogen mobilization from pod to seed
are depicted. The weight ratio of imports over the
life of the fruit is C : N : H 2 0 :: 12 : 1 : 600. (from
Pate et al. 1977; Sharkey 1977).

carbon, 89% of the nitrogen and 40% of the
water enters through phloem, the remainder
through xylem. A fully-illuminated fruit effects
net gains of carbon during the photo-periods of
all but the last two weeks of its life (Sharkey
1977), but in no week of its life do the net daily
gains of C0 2 from the outside atmosphere com-
pensate for the night time loss of C0 2 from the
fruit. Net respiratory losses increase dramati-
cally as the fruit ages, due to increasing C0 2
output from the seeds and loss of chlorophyll
and of photosynthetic performance by the pod.

Of every 100 units of carbon imported by a
white lupin fruit over its life, 52 are eventually
incorporated into seeds, 37 into non-mobilizable
materials of the pod, and the remaining 11 lost
as a net efflux of C0 2 to the outside atmosphere.
By contrast 96% of the nitrogen entering the
fruit ends up in the seeds, this element being
mobilised to the seeds from temporary storage
in the pod with 80% efficiency (Fig. 3).

The heavy dependence of legume fruits on
phloem-borne assimilates and their low attrac-
tion of xylem -mobile solutes through transpira-
tion, results in fruits, and especially seeds, having
a final composition closely resembling that of
phloem translocate (Pate et al. 1977; Hocking and

89

Journal of the Royal Society of Western Australia, Vol. 62, Parts 1-4, 1979.

Pate 1977). In particular they exhibit low levels
of elements only sparingly mobile in phloem
(e.g. Ca and Mn), and under conditions of short-
age of these elements seeds are likely to be the
first to show deficiency symptoms. This is well
displayed for manganese deficiency in the “marsh
spot” disorder of peas (Reynolds 1955) and for
the split-seed syndrome associated with defici-
ency of the same element in lupins (Perry and
Gartrell 1976; Hocking et al. 1977). Similarly a
reduction in transpiration of a fruit by enclosure
can cause deficiency symptoms of calcium to
occur in fruits, suggesting how critically impor-
tant the transpirational attraction of xylem-
mobile nutrients is in the normal development of
the fruit (see Wiersum 1966). In underground
fruits of legumes such as peanut the supply of
Ca through the fruit stalk is supplemented by
absorption of the element by the fruit surface
from the rooting medium (Bledsoe et al. 1949),
a property compensating for the meagre xylem
intake of the fruit.

Our studies of C0 2 fluxes between the pea
fruit and its external and internal atmospheres,
have enabled its photosynthetic performance to
be related to the overall carbon budget of the
fruit. During the first half of a 36-day growth
period (Fig. 4A; 0-18 days) much of the carbon
imported through the fruit stalk is consumed in
pod growth, and 14 C0 2 feeding studies show that
pod photosynthesis is to the benefit of the pod
rather than the contained seeds. At this time
the pod fixes more C0 2 from the external atmo-
sphere than from the pod gas space, largely
because the respiratory output of seeds at this
stage is small relative to the photosynthetic
potential of the pod. In the second half of fruit
development (18-36 days; Fig. 4B), pods have
ceased to accumulate dry matter so translocate
from the parent plant is directed entirely to
seeds. Seeds also benefit from pod photosyn-
thesis, especially the fixation of respired C0 2
from the gas cavity of the fruit.

As a result of pod photosynthesis a fully illu-
minated fruit of Pisum sativum requires over its
life 16-19% less carbon from the parent plant
than would a fruit laying down the same amount
of dry matter in total darkness (Flinn et al.
1977). It follows that agronomic practices
should be designed to maximise the efficiency of
light interception by fruits in the plant canopy.

From the viewpoint of seed production as a
nutritional source for animals the most signifi-
cant measure of fruit performance is how effici-
ently assimilates imported from the parent plant
are ultimately converted into digestible food
reserves of seeds. To make such estimates we
require knowledge of the weight of sugar and
nitrogenous solutes imported by the fruit during
its life, the final dry weight of the seeds, and the
proportion by weight therein of “useful” food
reserves such as starch, oil and protein.

A comparison between two species, white lupin
(Pate et al. 1977) and garden pea (Flinn et al.
1977), shows how grain legumes may vary in
these respects. Both of these species bleed from
their phloem and xylem (Pate et al. 1974), so the

14.7

15.5

i.'o

translocate

lllllllll photoperiod CO2 flux & photosynthate
HM | night CO2 flux
□ incorporation into dry matter
T ] net loss of dry matter

Figure 4. — Economy of carbon in developing fruits of
Pisum sativum L. (cv Greenfeast). Carbon fluxes are
shown in relation to intake of translocate and
exchanges of C0 2 between seeds and pod of the
internal and external atmospheres of the fruit. Day-
time respiration of the fruit is not shown. (A) 0-18
days after anthesis, a period when 351 mg C were
imported through the fruit stalk; (B) 18-36 days
after anthesis, a period when 797 mg C were imported
through the fruit stalk. The flux values in (A) and
(B) are expressed relative to an intake of 100 units
of carbon through the fruit stalk. (Data from Flinn
et al. 1977).

Identity of the translocated solutes received by
the fruit can be accurately assessed. This in-
formation, matched against the economy of
carbon, nitrogen and water of the fruit, allows
one to compute the precise weights of amides,
amino acids and sugars imported by the fruit
during its life. As shown in the data of Table
4 the pea fruit turns out to be the more efficient
converter of assimilates into seed dry matter
(69% in peas, 50% in lupin), or “useful” food

90

Journal of the Royal Society of Western Australia, Vol. 62, Parts 1-4, 1979.

substances (45% in peas versus 31% in lupin),
but the lupin fruit emerges as the superior
species if efficiency is measured in terms of pro-
duction of seed protein.

Table 4.

Efficiency of conversion of translocate into seed products
by fruits of two species of grain legume

Pisum
sativum
L. (cv

Greenfeast)

Lupinus
albus L.
(cv Neutra)

(A) Solutes translocated to fruits

(mg . plant - 1 )

sucrose

2 404

1 756

amides + amino acids

393

384

(B) Materials laid down in fruits

(mg . plant - 1 )

total dry matter

1 930

1 070

protein

387

412

oil

19

132

perchloric acid-soluble carbo-

hydrate

857

110

(C) Conversion efficiency by
weight of imported solutes to
fruit components (%).

to dry matter ....

69

50

to protein

14

19

to protein + oil + digestible
carbohydrate

45

31

Net photosynthate Total C gain as dry

(net C gain by = matter by plant day
shoot in day) and night

It remains to be seen how the values obtained
by the two fruits described above compare with
grain legumes as a whole, and whether better
conversion efficiencies might be achieved in fruits
by careful selection or breeding.

The carbon economy of whole legumes

Before judgment can be passed on the relative
efficiencies with which grain legumes operate as
crop plants, assessments must be made of their
total performance, especially in the context of
utilization of photosynthate, for nitrogen fixation
and, ultimately, for seed production. With this
in mind we have attempted to construct for
several grain legumes budgets for carbon utiliza-
tion during growth of the whole plant and its
parts, utilizing information on carbon gains or
losses as dry matter of plant parts, knowledge of
the compounds involved in root-shoot inter-
change via xylem and phloem, and assessments
of respiratory and photosynthetic exchanges of
C0 2 by plant organs over specific intervals of
the growth cycle. The budgeting takes as its
centrepoint the partitioning of the net photo-
synthate produced by photosynthetic surfaces of
the plant during the day, this quantity being
determined indirectly as: —

Respiratory loss Respiratory

-f of C from + loss of C by

shoot at night nodulated root

(day + night)

C

A

Phase 1 , vegetative (0-49 days )
net photosynthesis 2-38 gC
N ? fixation 0 116 g N

B

Phase 2, flowering and early fruiting (50-94 days)
net photosynthesis 24 30g C
N ? fixation 0-672 g N

Phase 3. seed filling (95-135 days)

net photosynthesis 14-77 g C
N ? fixation 0 293 g N

phloem- transported C as photosynthate
or as mobilised carbohydrate

rylem transported C with fixed N

KEY

□

0

C incorporated into dry matter
respired C loss as CO 2 1

.. phloem -reyc led C with mobilised N

C loss in abscised organs
C loss from dry matter

Figure 5. — The economy of carbon in nodulated plants of Lupinus albus L. (cv Neutra), relying solely on nodules
for their nitrogen supply. The situation in three phases of the growth cycle is described. In each case items
of the carbon budget are expressed relative to a net intake of 100 units of carbon by photosynthetic surfaces
of the shoot. The amounts of carbon and nitrogen assimilated during each phase are indicated. (Data from
Pate and Herridge 1978).

91

Journal of the Royal Society of Western Australia, Vol. 62, Parts 1-4, 1979.

Phase 1. vegetative (0-61d)

net photosynthesis 9 16 g C
N fixation 0 393 g N

stem

petioles

★

100

leaflets

nodules

B

Phase 2. flowering and early fruiting (62-78 d)
net photosynthesis 10-79 g C
N 2 fixation 0 333 g N

3*

reproductive parts

nodules

100

Phase 3. seed-filling (79-120d)

net photosynthesis 4 76 g C
N 2 fixation 0 065 g N

12 *

reproductive parts

fallen

petioles x - .

f 32; \

f 2 VJ i

if

C 2

roots

nodules

y

shed Q
nodules

12
28

CD

shed roots 40

* phloem- transported C as photosynthate
or as mobilised carbohydrate

— — xylem- transported C with fixed N

KEY

□ C incorporated into dry matter 0
respired C loss as CO2 £1)

..4.. phloem -reycled C with mobilised N

C loss in abscised organs
C loss from dry matter

Figure 6. — The economy of carbon in nodulated plants of Vigna unguiculata (L.) Walp. (cv Caloona) relying solely
on nodules for their nitrogen supply. The situation in three phases of the growth cycle is described. In each case
items of the carbon budget are expressed relative to a net intake of 100 units of carbon by photosynthetic
surfaces of the shoot. The amounts of carbon and nitrogen assimilated during each phase are indicated.
(Data from Herridge and Pate 1977).

By measuring respiratory losses and dry weight
changes in nodules and roots separately from
those of shoots, the significance of the plant’s
nodules as consumers of net photosynthate can
be assessed. Also, as described in an earlier
section on nodule functioning, the C:N ratio of
xylem exudate can be related to nitrogen incre-
ments of the shoot to estimate how much of the
carbon donated to the root, as photosynthate,
returns back to the shoot attached to products of
nitrogen fixation.

Information is available at present for only
two legumes, white lupin, a tap-rooted species,
grown as a winter annual in Western Australia,
and cowpea, a slender-rooted tropical legume
and a summer legume as far as culture in West-
ern Australia is concerned. The cultivars of both
species used for the experiments were character-
ized by low harvest indices for seed production,
and hence may be regarded as typical of the
many legumes needing selection for improved
yield of grain.

The carbon economy of these species is sum-
marized in Figures 5 and 6, the comparisons
relating to each of three stages of growth,
namely vegetative development prior to flower-
ing, flowering and early fruiting, and seed filling.
Each budget expresses items of expenditure of

carbon relative to a net intake of 100 units of
carbon as net photosynthate by foliar surfaces of
the plant. Attached to each budget is informa-
tion on the amounts of nitrogen fixed and net
photosynthate generated, so that the relative
importance of each phase of growth is depicted
relative to the total assimilatory output of the
plant during its life. Comments relating to each
phase of development are as follows: —

Vegetative growth . — Although the vegetative
phase of lupin lasted longer and encompassed a
much greater proportion of the plant’s growth
and assimilation than in cowpea, the species
show very similar patterns of partitioning of net
photosynthate. The largest single item is carbon
translocated to roots (54% of net photosynthate
in lupin, 41% in cowpea), most of this being
respired, some contributing to growth of root
and nodule, and a relatively small amount re-
turning to the shoot associated with products of
nitrogen fixation.

Flowering and early fruiting . — Both species
show greatest photosynthetic and nitrogen-fixing
capacities at this time. Lupin continues to
acquire almost half (49%) of the plant’s net
photosynthate, versus one-third (34%) in cow-
pea. The high consumption of carbon by roots
of lupin relates particularly to the establishment

92

Journal of the Royal Society of Western Australia, Vol. 62, Parts 1-4, 1979.

and maintenance of an extensive tap-root.
Whilst most desirable for growth in the dry,
nutrient-poor sands of south-west Australia,
this feature clearly carries penalties in terms of
decreased diversion of photosynthate to grain.

The proportions of photosynthate consumed by
nodules during flowering and early fruiting is less
than in vegetative growth, reflecting the ever
increasing sink capacity of shoots for photosyn-
thate. Four times more carbon is bound into
shoots than into nodulated roots in lupin, seven
times more in the case of cowpea, so the shoot:
root weight ratio widens progressively after
flowering.

Seed filling . — Most marked differences occur
between the species at this stage. The roots
share of photosynthate remains high (54%) in
lupin, but is much less (14%) in cowpea. As
might be expected, returns of fixed N from nod-
ules are lower during this phase in cowpea than
in lupin, and, due to a lower expenditure of car-
bon on roots, cowpea is able to divert relatively
more photosynthate to fruits than in lupin.
However, when measured in terms of efficiency
of mobilization of nitrogen to seeds, lupin proves
superior, 61% of its fixed nitrogen ending up in
seeds compared with only 43% in cowpea. This
is because the high protein seeds of lupin incor-
porate relatively more nitrogen than do the low
protein seeds of cowpea (see Tables 1 and 5).

Since the value of legumes stems mainly from
the protein which they synthesize, the efficiency
with which photosynthate is used for this pur-
pose is a very relevant measure of their func-
tioning as crop plants. The cultivars of cow-
pea and lupin examined in our investigations are
used in agriculture as green forage crops as well
as for grain production, so conversion efficiencies
are appropriate if based on photosynthate con-
sumption in relation to protein synthesized in
above-ground vegetative parts by the time of
flowering, or on the basis of protein harvested in
seeds at the end of the growth cycle. The rele-
vant calculations are shown in Table 5, where
it can be seen that 17-25 g net photosynthate
(as carbohydrate) are required to make each
gram of protein accumulated in above ground
parts by the time of flowering, 31-33 g carbo-
hydrate for each gram of protein harvested as
seed. Thus there is a conversion by weight of
only 4-5.8% of the net photosynthate produced
in vegetative growth into forage protein, an
efficiency of only 3% for the comparable con-
version to seed protein. It would be especially
interesting to see how the values for the rela-
tively unimproved cultivars studied here com-
pare with the situation in cultivars and species
of grain legumes selected for high yield of ^eed.
Such comparisons are in progress.

It is now appropriate to return to the original
question of how grain legumes compare with
other seed-yielding crops in productivity and
overall efficiency of performance. There simply
is not yet enough data available for such com-
parisons to be made, since the economy of carbon
in non-legumes has yet to be studied on strictly
comparable grounds to our own studies of
legumes, and it would indeed be most unwise to
attempt to generalize about legume performance

Table 5.

Costs in terms of net -photosynthate of protein
production by two grain legumes i 1 )

Vigna

unguiculata
(L) Walp.
(cv Caloona)

Lupinus
albus L.
(cv Neutra)

(A) Synthesis of protein in above-
ground vegetative parts ( 2 ).

Production of net photosyn-
thate (g carbohydrate
plant* 1 )

50.1

66.7

Protein accumulated in shoot
(g. plant* 1 )

2.91

2.70

Net photosynthate consumed
per unit of protein synthe-
sized (g carbohydrate . g
protein* 1 )

17.2

24.7

(B) Synthesis of seed protein < 3 >.
Production of net photosyn-
thate (g carbohydrate
plant* 1 )

61.8

103.6

Protein accumulated in seed
(g . plant* 1 )

1.90

3.34

Net photosynthate consumed
per unit of seed protein
synthesized (g carbohydrate
. g protein* 1 )

32.5

31.0

(1) Data for cowpea from Herridge and Pate (1977) and
for white lupin from Pate and Herridge (1977).

(2) Measured over the period from germination to the
time of maximum nitrogen content in above-ground
vegetative parts (i.e. at 79 days after germination in
Vigna, and at 95 days in Lupinus).

(3) Production of net photosynthate calculated for com-
plete growth cycle, seed protein measured as amount
present per plant at full maturity (i.e. at 120 days
in cowpea, and at 135 days in white lupin).

on the basis of investigations of only two species.
It would seem appropriate in further studies on
legumes to examine the mechanisms underlying
the substantial respiratory losses which occur
from roots and the processes whereby fruits and
nitrogen-fixing nodules compete for a dwindling
supply of assimilates at the end of the growth
cycle. Our data suggest that nodules, and their
fixation activities, are not particularly large con-
sumers of photosynthate, so that any inadequa-
cies which legumes may show in comparison
with non-legumes are likely to relate to the
overall performance of the host plant rather
than to the symbiotic fixation of nitrogen.

Acknowledgments. — The work described in this paper
has involved many persons whom I wish to thank
sincerely for the stimulation and companionship which
they have so freely given. Names of those collaborating
on various projects are to be found in the list of
references, but I would wish to acknowledge also the
technical assistance in Belfast of James Daly and Leah
Green, and in Western Australia the assistance of Ian
Passmore, Edvin Rasins, Kathy Hamel, David Waldie and
Glen Oakley. The work has been supported by the
Agricultural Research Council (U.K.), and Science Re-
search Council (U.K.), and by the Australian Research
Grants Committee, the Australian Meat Research
Committee, and the Wheat Industry Research Council.

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Perry, M. W. and Gartrell, J. W. (1976). — Lupin “split
seed” — a disorder of seed production in
sweet, narrow-leafed lupins. J. Agric. W.
Aust. 17: 20-25.

Reynolds, J. D. (1955). — Marsh spot of peas: a review of
present knowledge. J. Sci. Fd. Agric. 6:
725-734.

Schubert, K. R. and Evans, H. J. (1976). — Hydrogen
evolution: a major factor affecting the effici-
ency of nitrogen fixation in nodulated sym-
bionts. Proc. Nat. Acad. Sci. USA. 73: 1207-
1211 .

Schubert, K. R., Engelke, J. A., Russell, S. A. and Evans
H. J. (1977). — Hydrogen reactions of nodu-
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60: 651-654.

Sharkey, P. J. (1977). — Transport to and nutrition of the
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Siegel, A. and Fawcett, B. (1976). — Food legume process-
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Wallace, W. and Pate, J. S. (1965). — Nitrate reductase in
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Wiersum, L. K. (1966). — Calcium content of fruits and
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Journal of the Royal Society of Western Australia, Vol. 62, Parts 1-4, 1979, p. 95-107.

Aspects of Science in Western Australia 1829-1979.

A volume to commemorate the 150th Anniversary of the founding of Western Australia.

Studies on the resin chemistry of some
Western Australian plants

by P. R. Jefferies

Organic Chemistry Department, University of Western Australia, Nedlands, W.A. 6009

Abstract

Chemical research on plant resins in Western Australia is reviewed with
particular reference to components of the sub-family Ricinocarpoideae of the
Euphorbiaceae. The diterpenoids are discussed in terms of their relationships with
the gibberellin pathway. Siparate correlations between the bicarbocyclic and tstra-
cyclic groups are also given and in vivo experiments to define some of these relation-
ships are described. Microbiological hydroxylation of tetracyclic systems is also
summarised and an account of some experiments relating to gibberellin biosynthesis
is presented. The occurrence of triterpene and flavonoid components associated with
the resins is also given.

Introduction

As casual observers the late Professor White
and I were particularly impressed by the fre-
quency plant species in the Eremean and Wheat-
belt areas of Western Australia exhibit resin
coatings on their leaves and terminal branches.
The organic chemist is aware not only of indus-
trial applications of resins and allied substances
but also of the fact that these materials have
been one of the main sources of the polyterpenes.
In particular the most abundant of the resins,
those from pines, revealed the field of diterpenes
to organic chemistry. At the outset of this work
diterpenes were a small group of compounds
which were largely confined to the pines and so
we were interested to discover if these common
coatings were a major extension of this group.
As with the pine resins we were at first restricted
by the problem of separating what were often
complex mixtures and progress was accelerated
by sampling on a wide scale and selecting the
more tractable mixtures. Developments in
analytical and preparative methods of separation
allowed us to return later to the more complex
cases which are commonly encountered, particu-
larly in Eremophila. The project we began in
this area is a continuing one but those aspects
of it related to the sub -family Ricinocarpoideae
of the Euphorbiaceae have reached the stage
where they can be usefully reviewed. In view
of developing interest by biologists in plant resins
(Dell and McComb 1974, 1975) the opportunity is
also taken to catalogue resin components from
other local species. In the early stages the main
problems lay in the proof of structure and stereo-
chemistry. Advances in physical methods make
these problems secondary and since they are of
interest only to chemists this aspect of the work
is treated briefly. On the other hand the bio-
synthesis of many of these substances is of
general interest and in the sequel the structural

patterns are identified and correlated through
direct pathways although the existence of meta-
bolic networks is most probable. A number of
in vivo experiments have been carried out to
define more obscure biosynthesis and these are
discussed. The relationship of the ent- kaurene
diterpenes to gibberellins is of particular import-
ance and our contribution in this area is also
summarised.

Resins from species of Ricinocarpoideae

The first material chosen came from the wide-
spread Beyeria drummondii which gave ~20%
of ether soluble resin of which almost a quarter
was the cinnamate of a diterpenetriol with a new
carbon skeleton which we called beyerane. It
was clear that an extension to other members
of the family was warranted and wide-ranging
plant collections were undertaken to secure
material. These provided many of the known
species of Ricinocarpoideae along with a number
of new members some of which were ult ; mately
described at Kew (Airy Shaw 1971) through the
agency of the Western Australian Herbarium.
These included Beyeria calycina and B. calycina
var. minor, B. brevifolia var. truncata and var.
brevipes, and Bertya cupressoidea. Inevitably
many of our early papers used undefined names
and the relationship of these plants to the pre-
sent taxonomy has been summarised recently
(Errington et al. 1976). Most species of Beyeria
and Bertya carried resin, exceptions being the
coastal Beyeria cyanescens which lacked signifi-
cant coating and the near-coastal B. gardneri
which had very little. On the other hand B.
viscosa from Rottnest Island had a substantial
resin layer. No resin was observed on Monotaxis
spp. or on Ricinocarpus tuberculatus or R. vein-
tinus although the latter showed some toxicity
which was associated with low-level anti-tumor
activity. Amperea spp. have not been studied.

95

Journal of the Royal Society of Western Australia, Vol. 62, Parts 1-4, 1979.

In general the resins were hard materials
which consisted largely of widely varying propor-
tions of flavonoids, diterpenes and fatty material
with triterpenes appearing in Beyeria brevifolia,
B. viscosa and B. leschenaultii. The resin of B.
brevifolia var. truncata represents one extreme

in that it is essentially flavonoid whereas B. vis-
cosa and B. lepidopetala show no significant
proportion of these compounds. A list of the
main polyterpenoids and flavones isolated
together with salient references is given in
Table I.

Table 1

Chemical components of resins from species in the Ricinocarpoideae

Species

Polyterpenes (Structures) Flavones* References

Beyeria brevifolia (Muell. Arg.)
Benth var. brevifolia Airy Shaw

secopimaradiene (21),
lup-20(29)-ene-3j8, 1 6yg-diol, and
lup-20(29)-ene-3^3. 1 6^8.28-triol

5,7,3'-trihydroxy-3.8.4',5'-

tetramethoxy-

Chow and Jefferies 1968
Errington et al. 1976

B. brevifolia (Muell. Arg.) Benth
var. brevipes Airy Shaw

kauranes (59), (40)

Baddeley et al. 1964c

B. brevifolia (Muell. Arg.) Benth
var. truncata Airy Shaw

5-hydroxy-7,4'-dimethoxy-

5,4'-dihydroxy-7-methoxy-

5,7-dihydroxy-3,8,4'-trimethoxy-

Dawson et al. 1965

B. brevifolia (Muell. Arg.) Benth
var. nov.

secopimaradiene (21).
lup-20(29)-ene-3|8, 16^-diol and
lup-20(29)-ene-3^J, 1 6jg,28-triol.

5,7,4'-trihydroxy-3,8-dimethoxy-

5.7,4'-trihydroxy-3,8,3'-

trimethoxy-

5,7-dihydroxy-3,8,3',4',5'-

pentamethoxy-

Stacey 1970

B. calycina Airy Shaw

labdanes (10) (11)(12/(13)
kauranes (8)(39)(41)(42)(45)(46)

5,4'-dihydroxy-3,7-dimethoxy-

Jefferies and Payne 1965

B. calycina Airy Shaw var. minor
Airy Shaw

beyeranes (23)(25)(27)(29)
secobeyeranes (24)(26)(30)

Ghisalberti and Jefferies 1968,
Bakker et al. 1972

B. drummondii Muell. Arg.

beyerol (25)

5,4'-dihydroxy-7-methoxy-

5,7-dihydroxy-4'-methoxy-

5,7,4'-trihydroxy-

Jefferies et al. 1963
Stacey 1970

O’Connell and Maslen 1966

B. drummondii Muell. Arg. var. nov.

beyeranes (22)(25)(28)

5,3'.4'-trihydroxy-7-methoxy-

5,3'-dihydroxy-3,7.4'-trimethoxy-

5,7-dihydroxy-3,7,8-trimethoxy-

Ratajczak 1969
Jefferies unpublished

B. latifolia (Muell. Arg.) Baill.

kauranes (35)(38)

5,4'-dihydroxy-7-methoxy-

5,7,4'-trihydroxy-

Jefferies and Retallack 1968a

B. lepidopetala F. Muell.

labdanes (15)(16)(17)

Coates 1966

B. leschenaultii (D.C.) Baill.

kauranes (31)(35)(38)(43)(46)(48)
lup-20(29)-ene-3^3,16^3-diol

5,4'-dihydroxy-3,7,8-trimethoxy-

Jefferies and Retallack 1968b
Baddeley et al. 1964a,b

B. viscosa (Labill.) Miq.

kauranes (3 1)(32)(33)(34)(35)(36)
(37)(42)

lup-20(29)-ene-3^3, 16^S-diol
lup-20(29)-en-3-one

Coates 1966

Bertya cupressoidea Airy Shaw

bertyadionol group (50)(51)

Ghisalberti et al. 1974
Maslen et al. 1975

B. dimer ostigma F. Muell.

5,3',5'-trihydroxy-3,7,4'-trimeth-

oxy-

Henrick and Jefferies 1964b

Ricinocarpus muricatus Muell. Arg.

labdanes (14)(15)(16)(17)(18)(19)

5,7,3',4'-tetrahydroxy-3,8-dime-

thoxy-

5,3',4'-trihydroxy-3,7.8,-trimeth-

oxy-

5,3'-dihydroxy-3,7,8,4'-tetra-

methoxy-

Henrick and Jefferies 1965a.
1965c

R. psilocladus (Meull. Arg.) Benth.

5,7,3',4'-tetrahydroxy-

Stacey 1970

R. stylosus Diels

polyalthic acid (20 ) ;
kauranes (8)(4 1)(43)(44)(45)(47)

5,4'-dihydroxy-3,7,8-trimethoxy-

5,3'-dihydroxy-3,7.8,4'-tetra-

methoxy-

5-hydroxy-3,7,8,3',4'-penta-

methoxy-

Henrick and Jefferies 1964a
1964b, 1965b

* For numbering see Figure 8b.

96

Journal of the Royal Society of Western Australia, Vol. 62, Parts 1-4, 1979.

a. 1 up-20 (29) -ene-36-ol b. flavone

Figure 1. — A.- lup-20(29)-ene-3^-ol. B.- flavone.

Beyeria brevifolia, B. viscosa and B. leschen-
aultii provided large quantities of the triterpene
lup-20(29)ene-3£, 16/3-diol, which in the latter
plant was associated with a small proportion of
the 3/3, 16/3, 28-triol. Despite the fact that

lup-20(29)-ene-3/3-ol (Fig. la) and the 3/3,28-diol
are very widely distributed in plants neither the
3^,16-diol or the triol had been isolated pre-
viously. Evidently 16/3-hydroxylation of the
3/3-ol is an abberration of the hydroxylating
system deflecting attack from C-28 to the neigh-
bouring C-16. The structures of these triter-
penes were proved by interrelation with each
other and with known lupane derivatives.

Despite the fact that the flavones (Fig lb)
are widely distributed and well documented most
of the substances we isolated were new mem-
bers of the class. The resin flavonoids are excep-
tional in that not only are they highly methoxy-
lated but they frequently occur as a large pro-
portion of the plant material. A number of
common structural features were observed and
these include the ubiquitous 5 -hydroxyl group
and the fact that when substitution occurs at
C-3 or C-8 then the substituent is always
methoxyl. Oxygenation of the A ring follows
the normal 5,7 pattern, established by assembly

of malonyl CoA residues and later substitution
is limited to C-8; the common alternative C-6
is not substituted. Substitution of the phenyl
ring follows the normal patterns 4’; 3’, 4’; 3’,
4', 5’. In non-resin flavones methylation of B
ring hydroxyl groups occurs almost exclusively
for the 3 ’-OH and an enzyme system effecting
this reaction has been characterized (Ebell et al.
1972). No such specificity is observed in the
resins and both the 3’ and 4’-OH frequently show
methylation. Many roles have been suggested
for flavonoids in plants (McClure 1975) and
some of these functions no doubt apply in the
resins. Thus the ability to absorb UV light
enhances the protective nature of the coating.
They may also protect the resin from microbial
attack and as well, act as antioxidants to stab-
ilise the resin.

In considering the chemistry of the diterpenoid
constituents it is useful to relate them to the
biosynthesis of this group (Fig. 2). It is now
well established (e.g. Hanson 1972) that diter-
penes along with the steroids and triterpenes
arise from acetate through mevalonate leading
to the C 5 pyrophosphates which provide geranyl
pyrophosphate. Successive elimination-addition
reactions of two isopentenyl pyrophosphate mole-
cules then gives geranyl geranyl pyrophosphate
which with some stereochemical limitations may
be considered to follow divergent paths to macro-
cyclic ring systems such as casbene and the cem-
brenes, undergo cyclisation of the three terminal
C 5 residues to give C 2 o analogues of the sesquiter-
penes, or the all trans isomer may undergo
cyclisation to give the bicyclic labdane ion (1)*.
Backbone rearrangement of ( 1 ) gives members of
the eni-clerodane group whereas deprotonation
provides copalol pyrophosphate (2). The latter
may cyclise first to the tricyclic pimarene (3)

7 , R - CI^OH ) _8 , R = CO 2 H > Gibberellins

Figure 2. — Scheme for the biosynthesis of some en£-diterpenes.

* Numbers in parenthesis refer to structures in the figures.

97

82710 ( 7 )

Journal of the Royal Society of Western Australia, Vol. 62, Parts 1-4, 1979.

and then to the tetracyclic beyerane ( 4 ) and
kaurane ions (5). The pathway is drawn to
show free carbonium ions and although such
species have not been proved to arise in biological
systems they represent a convenient basis for
rationalising the structural assembly. Ent-
kaurene (6) is converted by way of the 19-
hydroxy derivative (7) to the kaurenoic acid
(8) which undergoes 7/3-hydroxylation before
generating gibberellins. In view of the wide-
spread distribution of these latter compounds
and their physiological role, the pathway to gib-
berellins may be regarded as part of primary
metabolism in plants. The section from the
bicyclic carbonium ion pyrophosphate (I) to
kaurenoic acid ( 8 ) is a useful framework for
discussing the diterpenes of Ricinocarpoideae
since with the exception of the bertyadionol
group, which are present in Bertya cuppressoidea,
all can be regarded as deriving from intermedi-
ates on this pathway. The diterpenes of many
other species are similarly derived. It is noted
specifically that the stereochemistry of C-10 in
this series is enantiomeric to that in the steroids,
triterpenes and common pine resin acids. For
the diterpenes, divergence occurs for the cyclisa-
tion of geranyl geranyl pyrophosphate which can
give either the bicarbocyclic labdane (I) or its
mirror image.

Bicarbocyclic diterpenes

The bicarbocyclic ion (1) is the parent of a
wide range of substances which were obtained
and rational pathways linking them, together
with their distribution among the species are
shown in Fig. 3. Hydration of 1 and pyrophos-
phate hydrolysis gives the diol (10) and this or
the pyrophosphate are the logical precursors of
the epimanoyl oxide (11) its 18-hydroxy (12)
and carboxy derivatives (13) all four of which
are found in B. calycina. The presence of a reduc-
tase generating the 13S configuration appears in
both B. lepidopetala and Ricinocarpus muricatus
leading from 10 to the diol (14) present in the
latter, and then by hydroxylation of C-18 to the
triol (15) found in both. Low specificity is indi-
cated for the 13S reductase since both plants give
13S systems which arise from 9. Thus reduction
of 9 and hydroxylation and oxidation of C-18
or C-18 and C-15 give the carboxylic acids (16)
and (17) respectively. Further hydroxylation of
the hydroxy acid (16) occurs at C-16 to give
the dihydroxyacid (18) found in R. muricatus.
Polyalthic acid (20) in R. stylosus and the but-
enolide (19) in R. muricatus must arise by
hydroxylation of C-16 at the 13-ene level coupled
with further oxidation of the C-15 alcohol to
the aldehyde in the former case and to the acid

Figure 3. — Probable biosynthetic relationships of the bicarbocyclic group.

98

Journal of the Royal Society of Western Australia, Vol. 62, Parts 1-4, 1979.

in the latter. The intermediate enediol group
has been observed in diterpenes from Sciadopitys
(Sumimoto et al. 1964). The hydroxylation and
oxidation of C-18 in (19) and (20) might occur
at any stage.

The chemical proof of the skeleton of these
compounds is based ultimately on chemical inter-
conversion with each other and with materials
of known constitution, and positions of the
hydroxyl and carboxyl groups follow largely
from spectroscopic and pK measurements. The
main ambiguity which required resolution was
the configuration of the C-13 position. This was
shown to correspond to that of eperuic acid
which was however unsettled. We were able to
clarify this situation (Henrick and Jefferies
1965a) since compounds with the 13R configura-
tion were available to us from simultaneous re-
search on Dodonaea lobulata resin (Dawson
et al. 1966). That these were epimeric at C-13
with those from Ricinocarpus followed from the
laboratory preparation of both from the diol
(10). Within the various groups of polycyclic
diterpenes oxygenation occurs commonly at
either C-18 or C-19. The bicarbocyclic examples
listed above show oxygenation at C-18 and not
C-19, in sharp contrast with the tetracyclics
described below, which exhibit oxygenation of
C-19.

Figure 4. — Origin of the seco-pimaradiene (21).

Tricarbocyclic diterpenes

The families of tricarbocyclic diterpenes derive
from a stereoisomer of the intermediate ion
formally shown as 3. Beyeria brevifolia var.
brevifolia and an undescribed variety both
generate compounds based on this skeleton. The
resin of the former contains mainly one diter-
pene (21, Fig 4) obtained as ~5% of the leaves
and branchlets. The efficient production of 21
is noteworthy since evidence presented in the
sequel suggests that it is apparently at least four
steps from the main path. Thus after formation
of the 8-ene from 3, hydroxylation of both C-3
and C-19 are evidently required before frag-
mentation of the A-ring can occur. Analogous
A-ring splitting is seen in Beyeria calycina var.
minor but has not been reported for other diter-
penes although several examples are known in
the triterpenes. Hydroxylation of the vinyl
group could occur at any stage and is analogous
to the 16,17-dihydroxylation of kaurene discussed
below. The structure assigned to 21 follows from
its interconversion with a derivative of beyerol.

Beyeranes

The diterpenetriol (25, Fig. 5) from B. drum-
mondii proved to have a new carbon skeleton
which was called beyerane and was recognised
as the missing link (4, Fig. 2) in the formal
pathway from the tricyclic ion (3) to kaurene.
The structure was proved largely by classical
oxidative degradation and dehydrogenation with
the limited application of physical methods and
the absolute stereochemistry was shown to be
antipodal to the steroids by optical rotatory dis-
persion methods. Small quantities of the diols
(22) and (28) were also isolated from a variant.

ent- beyer-15-ene , R,R ,R =H

28 R=H ; R'R"=OH

29

Figure 5. — Probable relationships between ent - beyerene metabolites.

30

99

Journal of the Royal Society of Western Australia, Vol. 62, Parts 1-4, 1979.

Beyeria calycina var. minor provided a group
of beyerene metabolites of which the more
abundant are the 6-acetate of the ketodiol (23)
and the seco-acid (26) which has undergone
A-ring fission analogous to the pimaradiene (21).
Several methods of effecting this ring splitting
have been developed in the laboratory. Thus
photolysis (Arigoni et al. 1960) of 3-ketones, as
for example 23, gives rise to the acids (24)
bearing an isopropyl group as an A-ring frag-
ment. Acids in which this residue appears as
isopropenyl (26) can be obtained by several
routes including the abnormal Beckman rear-
rangement (Witham 1960) of oximes of 3-ketones
such as (23). Both these methods were used to
relate beyerol (25) to the seco-acid (26). A
biomimetic fragmentation based on the reduction
of 3-keto-19-tosylates was also developed (Ghis-
alberti and Jefferies 1968).

Several schemes had been proposed to account
for the fragmentation process in vivo although
no experimental evidence in living systems was
available. These schemes fell into groups either
requiring C-18 and C-19 as methyl groups or
those for which one of these was hydroxylated
with variations within the groups arising from
different oxidation states of C-3. It seemed
likely that some distinction between these mech-
anisms might be made by feeding both enf-beyer-
15-ene and en£-beyer-15-ene-19-ol (27) to intact
plants since 19-hydroxylation is unlikely to be
reversible. Seedlings of B. calycina var. minor
were readily available following a bushfire and
these were transplanted to a glasshouse where

rapid growth occurred with regular watering.
Resin production was not obviously inhibited
and the normal resin components were found to
be present. 3 H labelled samples of enf-beyer-
15-ene, ent-beyer-15-ene-19-ol, (27), and the
3a, 19-; 19,17- (28) and 3a,17-diols (22) were pre-
pared and fed to these plants. We observed
incorporation into 26 for all substrates, clearly
indicating that the 4,4-dimethyl group is not
essential for the fragmentation and that mech-
anisms based on 3,19-dioxygenated substrates
are more probable (Bakker et al. 1972, Sefton
1978).

Beyeria calycina var. minor also contains small
quantities of the seco-acids (24) and (30) which
could be obtained independently by ultra-violet
irradiation of the co-occurring 3-ketones (23)
and (29) respectively. The possibility that these
acids arise in the plant resin by lengthy exposure
to sunlight is supported by our observation that
during the short periods required for incorpora-
tion of radioactivity from the labelled substrates
into 26 no significant radioactivity was detected
in 24. The retention of configuration at C4 in
the sequence 29 30 although expected for a

bio-transformation was not predicted for the
photochemical reaction and is attributed to
restriction to rotation of the C4-C5 bond.

Kauranes

Many of the plants we examined were found
to contain metabolites of enf-kaurene (6). The
carbon skeletons of these substances were estab-

Figure 6. — Probable relationships between ent-kaurene metabolites.

100

Journal of the Royal Society of Western Australia, Vol. 62, Parts 1-4, 1979.

lished by interconversion with each other and
with the parent enf-16/3-kaurane whose structure
and stereochemistry had previously been resolved.
A major problem in effecting these interconver-
sions lay in the unsatisfactory deoxygenation
methods available, particularly for hindered
primary alcohols, and for this purpose we dev-
eloped a technique involving the formation of
benzyl thioethers from sulphonate esters using
dipolar aprotic solvents (Henrick and Jefferies
1964a). Desulphurisation of these ethers to give
the hydrocarbons proceeded smoothly.

A schematic relationship showing possible
pathways between the oxygenated kaurenes is
shown in Figure 6 together with the range of
compounds associated with each species. Branch-
ing from the gibberellin pathway occurs by
hydroxylation at C-3 of either enf-kaurene ( 6 )
or the kaurenol (7) but this is not observed for
the kaurenoic acid (5). It is noted that C-3 and
C-19 are similarly situated with respect to the
D ring polar alkene group and since it is evident
from microbiological hydroxylation of steroids
(Jones 1973) that the distances from the binding
sites control sites of hydroxylation it appears
that the C-3 oxygenase and kaurene C-19
oxygenase have similar spatial demands. The
C-3 and C-19 oxygenase activities operate inde-
pendently since R. stylosus and B. calycina are
deficient in the former and B. viscosa has the
latter inhibited whereas the other species exhibit
both about equally. The 3-and or 19-oxygen-
ated kaurenes apparently can then undergo an
unspecific hydroxylation of the double bond
leading to 16, 17- diols. If hydroxylation of
C-3 and C-19 is dictated by binding to the D
ring double bond and if it is assumed that a
binding site may in turn become a hydroxylating
site (Jones 1973) then the C-3 and the C-19
hydroxyls might direct D ring hydroxylation. In
the beyerenes this could explain C-17 hydroxy-
lation and in the kaurenes 16,17-dihydroxylation
can be rationalised similarly since it has been
shown in metabolism of steroids by certain
moulds that substitution of a saturated hydroxy-
lation site by a double bond can lead to an
epoxide (Bloom and Shull 1955) and the latter
can readily generate a 1,2-diol. The formation
of these 16,17-diols appears to be general and
serves incidentally to remove kaurenol (7) and
kaurenoic acid (8) from the gibberellin pathway.
An alternative fate for the 16-ene is the forma-
tion of either the C-17 hydroxy or carboxy deri-
vative, through steps which appear to be avail-
able to most plants examined and which permit
a wide choice of substrate. Evidence presented
below indicates that these substances derive from
sequences analogous to the pinacol rearrange-
ment and probably arise through the diol dir-
ectly or the intermediate epoxide. In either case
the aldehyde is the immediate product and this
can distribute itself betwen primary alcohol and
carboxylic acid. The major kaurene metabolites
are thus accounted for. Two minor components
are the la,19-dihydroxy-17-acid (44) which
co-occurs with the 19-hydroxy- 17-acid (43) in
R. stylosus and the 12/3,17-dihydroxy-19-acid
(48) which co-occurs with the 17-hydroxy-19-
acid (46) in B. leschenaultii. These products

can be rationalised as arising irom the third
binding site of the hydroxylating system. Thus
primary binding to the C-19 carboxyl with sec-
ondary binding to C-17 H 2 OR is assumed to
direct the third site to C-12/3. Primary binding
to C-17 carboxyl with secondary attachment
C-19 H 2 OH could then result in C-la hydroxy-
lation since the C-17 -» C-12 and C-l -> C-19
distances are similar.

In summary then in its simplest terms the
secondary metabolism process in Figure 5 can
be explained by the operation of one hydroxy-
lating system with three binding Hydroxylating
sites along with an isomerase responsible for the
rearrangement of the 16,17-diols leading to C-17
alcohols and acids.

The formation of acid and primary alcohol
functions at C-17 has little biological precedent
and warranted further study (Croft et al. 1978).
Five possibilities which may be considered are
shown in Figure 7. Mechanism (a), an anti-
Markovnikoff hydration has been postulated for
a similar conversion of enf-kaurene in barley
seeds (Murphy and Briggs 1973). Mechanisms
(bi) and (b 2 ) involve pinacolic rearrangements
either through the epoxide or its diol and have
strong chemical analogy. Mechanism (c) has its
analogue in the dioldehydrase reaction and per-
mits such exchange of Hi on the enzyme complex
(Frey and Abeles 1966). Mechanism (d) involves
a dehydratase reaction leading to an enol. The
mechanisms predict four different fates for the
vinyl protons in 8 during its transformation to

Figure 7. — Mechanisms for the biosynthesis of the
hydroxy acid ( 46 ) from ent-kaurenoic acid ( 8 ).

101

Journal of the Royal Society of Western Australia, Vol. 62, Parts 1-4, 1979.

46. To distinguish these mechanisms kaurenoic
acid labelled with 14 C and at C-17 was pre-
pared and fed to seedlings of B. calycina. The
results show that no tritium was lost thus exclud-
ing mechanisms (c) and (d), and that about half
of the tritium had migrated to C-16 consistent
only with mechanisms of type (b). Distinction
between routes (bi) and (b 2 ) has not been made.

Diterpenes of Bertya cuppressoidea

It has recently become clear that a number of
species of Thymeleaceae and Euphorbiaceae, par-
ticularly Euphorbia and Croton produce diter-
penes which do not derive from the bicyclic ion
1. Some of these are potent co-carcinogens, of
v/hich the esters of phorbol (54, Fig. 8) are
best known (Hecker and Schmidt 1974).

The pathway to the carbon skeleton of phorbol
has not been established but clues to a likely
route are provided on the one hand by the struc-
tures of the Bertya diterpenes (Ghisalberti et al.
1973a, 1974) and by the related lathyrol group
(Opferkuch and Hecker 1973) and on the other
by casbene (49). The latter is produced by an
enzyme preparation of Ricinus communis (Euph-
orbiaceae) acting on geranyl geranyl pyrophos-
phate (Robinson and West 1969). Bertyadionol

(51) is the most abundant diterpene of Bertya
cuppressoidea and co-occurs with diterpene-D
(50) and -B, the C9-ene. The structures were
assigned by interrelation with each other, coupled
with extensive spectroscopic studies and the
absolute configuration was established by oxida-
tion to R-methylsuccinic acid and to the cyclo-
propane derivative homocaronic acid. The
structural relationships between the Bertya
diterpenes strongly suggest that the five-mem-
bered ring results from aldol condensation of a
diketone, which could be formed by allylic oxida-
tion of casbene at C-8 and C-ll, resulting in a
C7-C11 bond. Oxidation of C-4 and C-12 would
then provide diterpene D and bertyadionol.
Diterpene B could arise similarly with retention
of the 9-ene after inversion to the Z configura-
tion. Further cyclisation of this skeleton to give
the phorbol ring system requires linking C-12
and C-3, perhaps by an aldol type cyclisation
promoted by the C-5 substituent at the aldehyde
level (Worth 1971).

Bertyadionol has anomalous UV absorption
suggesting interaction between the cyclopenten-
one and vinylcyclopropylenone chromophores.
Irradiation of bertyadionol with UV light readily
affords a photo product as a consequence of an
unprecedented sequence leading to structure

(52) (Ghisalberti et al. 1978a). The photopro-
duct (diterpene C) accompanies bertyadionol in
the plant extracts and is probably formed in the
surface resin by solar radiation. Photolysis of
diterpene-D (50) results in an isomer (53) in
which the 13-ene has undergone the unexcep-
tional E-Z isomerisation. The inversion of C-2
which also occurs is probably due to ring fission
generating a C-15 carbene, rotation of the
C-l:C-2 bond and recyclisation as indicated in
Figure 8.

Figure 8. — Relationships between the Bertya diterpenes.

Mould metabolism of kauranes and beyeranes

Formal analogy between the plant and mould
hydroxylating systems prompted us to carry out
a survey (Beilby et al. 1973) of the microbial
hydroxylation of a group of tetracyclics using the
micro-organisms Calonectria decora, Rhizopus
nigricans and Aspergillus ochraceous for which
much data had been obtained on steroids by the
Oxford group (Jones 1973). It was hoped on the
one hand that plant hydroxylation patterns
might be reproduced by appropriate mould-
substrate choice and on the other we expected
that the moulds in offering a wider range of
hydroxylation systems might provide easy access
to B and C ring hydroxylated derivatives which
were not readily available from natural sources.
As it turned out both expectations were realised
in part (Table 2). Thus naturally occurring
1-hydroxykauranes which may be precursors of
the skeleton of the grayanotoxins occur rarely
but are easily available from 16,19-dioxygenated
norkauranes using R. nigricans. Hydroxylation
at C-13 is widespread in plant gibberellins but
otherwise rare in natural kauranes. This step
can be effected with some 16,19-dioxygenated
substrates using A. ochraceous and although
the yield is low it is comparable to synthetic
procedures and much more convenient.

102

Journal of the Royal Society of Western Australia, Vol. 62, Parts 1-4, 1979.

Table 2

Metabolism of some kaurenes by moulds *t

Substrate

Substrate modifications

Calonectria decora

Rhizopus nigricans

Aspergillus ochraceous

e/tf-kaur-16-en-19-oic acid (8)

7 a -OH; 15 a -OH; 7 a , 15 a -diOH

7/3-OH; 16 a ,17-diOH

16 a ,17-diOH

oxygenated ent- 1 6-nor-kaur- 1 6-ones
i 3 a -OH

7 a -OH

la-OH; 7 a -OH

6/3-OH ; 7 a OH

ii 19-acid

7 a -OH; 7^-OH; l a -OH

7 a -OH; 7/3-OH; l a -OH

13-OH; 13,16/3-diOH

iii 19-OH

7 a -OH; la-OH

7 a -OH; l a -OH

7 a -OH; 1 6/3-OH

♦Beilby et al. 1973; Ghisalberti et al. 1977a. t For numbering see Figure 6.

Hydroxylation of the B ring of kaurene is pre-
requisite to gibberellin formation and although
no B ring oxygenated metabolites of kaurene
were observed in the resins 6-oxygenated beyer-
anes were isolated. A number of beyerane
derivatives underwent 6/3-hydroxylation using
C. decora and similarly this hydroxylation tech-
nique was used to establish the skeleton of the
dihydroxy acid (29) (Fig. 4) since the derived
triol could also be obtained from the metabolism
of the diol from the authentic beyerane (27)
(Sefton 1978). All three moulds effected 7/3-

and 7a-hydroxylation with a wide range of
kaurane derivatives and application of these and
similar methods provided a simple route to
7/3-hydroxykaurenoic acid (Croft et al. 1974),
the gibberellin precursor otherwise available
only by lengthy procedures. The hydroxylation
by C. decora of kaurenoic acid gives the 15a-
hydroxy derivative a process which also occurs
in the plant Phebalium rude which contains both
these substances. Another example of plant
hydroxylation effected by moulds is the forma-
tion of the 16,17-diol by A. ochraceous from
kaurenoic acid.

60

Figure 9. — Metabolism of some ent - kaurenes by Gibberella fujikurci.

103

62

Journal of the Royal Society of Western Australia, Vol. 62, Parts 1-4, 1979.

Ent-kaurenes as sources of gibberellins

When kaurenoic acid (8) was first obtained a
report had just appeared indicating that its
13 -hydroxy derivative, steviol showed gibberellin
activity (Ruddat et al. 1963) and accordingly a
joint programme was established with Dr. B. O.
Phinney to determine the extent of this bio-
activity within our group of kaurene metabolites.
Although it was found that the effect was not
general for the kaurane nucleus ent - kaurene
(6), enf-kaurenol (7) and enf-kaurenoic acid
(8) did behave as gibberellins in a group of
dwarf maize mutants (Katsumi et. al. 1964a,
1964b). Since enf-kaurene itself was known to
be a precursor of gibberellic acid it seemed likely
that these substances were not intrinsically
active but were converted to gibberellins during
assay. This proposition was supported by feed-
ing experiments in Gibberella fujikuroi by several
groups using material of Western Australian
origin (Graebe et al. 1965, Verbiscar et al. 1967).
The sequence enf-kaurene kaurenol -» kaur-
enal -» kaurenoic acid -> gibberellins

which emerged from this work was supported
by work with plant systems (Dennis and West
1967). The formation of gibberellins from ent-
kaurenoic acid (8, Fig 9) clearly involved oxida-
tion of ring B but despite some effort the appro-
priate metabolite had not been obtained and so a
number of kaurene derivatives were fed to G.
fujikuroi in the hope that the hydroxylation
patterns obtained would identify the next step.
The results implicated C-7/3-hydroxylation since
this occurred in all cases (Jefferies et al. 1970,
1974c). Thus metabolism of the succinate ester

(55) of ent -kaurenol gave the 7/3- and
6/3,7/3-hydroxylated esters (Fig. 9). This experi-
ment was repeated with the isotopically labelled
kaurenol ester (55) and the 7/3-hydroxy ester
formed was hydrolysed to give the 7/3,19-diol

(56) which was found to be a precursor of gib-
berellic acid (59) . Fnt-kaurene-6/3,7/3,19-triol

(57) which was also formed from the suc-
cinate (55) did not produce gibberellic acid
in the mould although it possesses gibberellin
activity. In the meantime other groups had
reached the same conclusions (Hanson et al.
1972, Cross et al. 1970).

One apparently unnatural kaurene which
shows activity in maize is the dienol (58) . When
metabolised by G. fujikuroi this substance gives
a series of C-19 and C-20 gibberellins indicating
pathways to gibberellin A 5 (61) and As (62)
(Bakker et al. 1974) neither of which are normal
mould metabolites but are limited naturally to
plants. Backfeeding experiments showed that
the epoxidation step was probably most active
for the aldehyde (60) level of C-7 which was
also implicated in the removal of C-20.

Resins of other species

A selection of the many species outside the
Euphorbiaceae which carry resin has also been
studied and the results, with salient references,
are summarised in Table 3. A number of plants
are included which appear to lack external resin
but either contain diterpenes in significant
quantity or belong to genera which do.

Several Acacia spp. are viscid and the diter-
penes of A. rossei have been characterised as
bicyclic compounds arising from rearrangement
of the ion (1). The presence of kaurenoic acid
(8) and its 15a-hydroxy derivative in Phebalium
rude is a rare case of the isolation of diterpenes
from members of Rutaceae. On the other hand
many Western Australian species in this family
produce eriostoic or eriostemoic acid (Duffield
et al. 1962) both of which resemble diterpene
acids in the possession of twenty carbon atoms,
a carboxyl group and a polycyclic skeleton. Per-
haps they fulfill similar roles.

Reference has already been made to labdanes
in Dodonaea lobulata and similar compounds
were found in D. alternifolia and D. ptarmici-
folia. Backbone migration in the ion 1 (Scheme
1) gives the ent - clerodane skeleton which is the
basic structure of the compounds from several
other Dodonaea spp. Resin is commonly seen
on some members of the Boraginaceae and of
these Halgania lavandulacea has a heavy coating
and was selected for study. The resin was found
to be essentially a mixture of simple flavones.

Within the Discrastylidaceae the decorative
Cyanostegia angustifolia was found to be an-
other source of enf-clerodanes. Pityrodia lepi-
dota has a similar component whose relative
configuration was deduced by X-ray methods.
The resin of Newcastelia viscida contains an
isopimara-9(ll),15-dienediol which is the only
example we have encountered which has been
shown to belong to the so called normal series
(C-10/3 CH 3 ) which corresponds to biosynthesis
through structures enantiomeric with those of
Figure 1. The stereochemistry of this compound
coupled with the 9 (11) -double bond suggests that
it might arise from anomalous folding of geranyl
geranyl pyrophosphate leading to a trans, syn
stereoisomer of ion 1.

Eremophila species are probably the most
abundant resin plants of the Eremean and have
been found to contain a great diversity of diter-
pene skeletons. The compounds we have isolated
do not evolve from the ion 1 but are either
macrocyclic cembranoids or are generated by
sesquiterpene type cyclisations involving the
three C-5 residues adjacent to the pyrophosphate
of a geranyl geranyl pyrophosphate. Although
most of the resins contain significant quantities
of fatty material these rarely dominate.
Mvoporum beckeri is exceptional in that its very
sticky coating is largely fatty acids and alcohols.
Eremophila oppositifolia has internal viscid
material which contains a branched triply un-
saturated fatty acid.

Two Goodenia spp. have also been examined
and both contain esters of the kaurene-3a,19-
diol (31) previously obtained from B. leschen-
aultii; indeed G. strophiolata is the best source
of 3,19-functionalised kaurenes. One member of
the Asteraceae which has been well studied is the
perennial Olearia muelleri whose resin is another
source of ent-clerodanes. The low level anti-
tumour activity of this plant is associated with
flavone components.

104

Journal of the Royal Society of Western Australia, Vol. 62, Parts 1-4, 1979.

Table 3

Chemical components of other resinous and allied species

Plant Species

Polyterpenes, Fatty Acids

Flavones (or Aromatics)

References

MIMOSACEAE—

Acacia rossei F. Muell.

(8. 1 0)-friedolabdanes

5-hydroxy-7,4'-dimethoxy-

Langley 1966

RUTACEAE—

Phebalium rude Bartl.

kaurenoic acid (8)

1 5a-hydroxykaurenoic acid

Cannon et al. 1966

SAPINDACEAE—

Dodonaea inaequifolia Turcz.

ewMabdanes

Payne unpublished

D. lobulata F. Muell.

ewMabdanes

5-hydroxy-3,6,7,4'-tetramethoxy

Dawson et al. 1966

D. ptarmicifolia Turcz.

enMabdanes

Payne unpublished

D. microzyga F. Muell.

e/iMabdanes

Jefferies et al. 1974a

D. attenuata A. Cunn.

e///-clerodanes ....

Ghisalberti et al. 1973b

D. attenuata A. Cunn. var.

ent- clerodanes, lupanes

5.7-dihydroxy-3,6,4'-trimethoxy-

Payne and Jefferies 1973

linearis Benth.

D. boroniae folia G. Don

e/i/-clerodanes, ....

5-hydroxy-3,6.7,4'-tetramethoxy-

Jefferies et al. 1973

BORAGINACEAE—

Halgania lavandulacea Endl.

5,4'-dihydroxy-7-methoxy-

5.7- dihydroxy-4'-methoxy-

5.7- dihydroxy-3,4-dimethoxy-
5-hydroxy-3,4',7-trimethoxy-

J Stacey 1970

DICRASTYLIDACEAE —

Cyanostegia angustifolia Turcz.

(?«/-clerodanes ....

5,4'-dihydroxy-3.7,8-trimethoxy-

5.7,4'-trihydroxy-3,8-dimethoxy-

5,7,4'-trihydroxy-3,8,3'-trim-

Ghisalberti et al. 1967
Jefferies et al. 1973

ethoxy-

C. microphylla S. Moore

5.4'-dihydroxy-3,6,7-trimethoxy-

5,3'.4'-trihydroxy-3,6,7-trim-

Ghisalberti et al. 1967

ethoxy-

Newcastelia viscida E. Pritzel

ursolic and betulic acids isopim-

5,4'-dihydroxy-3,6,7-trimethoxy-

Jefferies and Ratajczak 1973

aradienediol ( 21 )

Pityrodia lepidota (F. Muell.)

clerodanes

Ghisalberti et al. 1978b

E. Pritzel

MYOPORACEAE—

Eremophila alternifolia R.Br.

5.7- dihydroxy-3-methoxy-

3. 5.7- trihydroxy flavanone

Jefferies et al. 1962

E. clarkei F. Muell. R.Br

cembranes

Coates et al. 1977
Maslen et al. 1977a

E. decipiens Ostf.

decipianes

Ghisalberti et al. 1975
Maslen et al. 1976

E. drummondii F. Muell

serrulatanes

Croft 1977

E. fraseri F. Muell. (var.)

eremanes (cembranes) ....

5,3'5'-trihydroxy-3,6,7,4'-tetra-

methoxy-

Jefferies et al. 1962
Oh and Maslen 1968

E. georgei Diels (var.)

cembranes (eremane, preziza-

Ghisalberti et al. 1976, 1977b,

enes)

Carrol et al. 1976, Maslen et
al. 1977b

E. glabra R.Br. Ostf.

(lirioresinol-B dimethyl ether)

Jefferies et al. 1961

E.longifolia F. Muell.

(safrole, eugenol methyl ether)

Della and Jefferies 1961

E. oppositifolia R.Br

fatty acids

Jefferies and Knox 1961

E. ramosissima C.A. Gardn.

5,7-dihydroxy-3-methoxy-
3, 5, 7, -trihydroxy flavanone

Jefferies et al. 1962

E. serrulata (A. Cunn.) Druce

serrulatanes

Croft et al. 1977

Myoporum beckeri F. Muell.

fatty acids

5,3'.4'-trihydroxy-7-methoxy-

flavanone

Goh 1968

GOODENIACEAE —

Goodenia strophiolata F. Muell.

kaurenediol (31)

5,7,3'4'-tetrahydroxy-3-methoxy-

Middleton and Jefferies 1968

G. ramellii F. Muell

kaurenediol (31), labdane

Coates et al. 1968

ASTERACEAE—

Olearia muelleri (Sond.) Benth.

e/f /-clerodanes ....

5,7,3'-trihydroxy-3,6,4'-trim-

ethoxy-

5,7, 3'4'-tetrahydroxy-3, 7-dim-

Jefferies et al. 1974b

beyeren-19-ol (27)

ethoxy-

Helipterum craspedoides W. V.

Dennison and Mirrington 1975

Fitzg.

Acknowledgements . — It is a pleasure to acknowledge
the continuing help we have had from the staff of the
Western Australian Herbarium on all the botanical
aspects of this project including identification and
storage of voucher specimens. We are grateful to Dr.
J. W. Green who accompanied us on some early collect-
ing trips and taught us a lttle taxonomy. My colleagues
Dr. J. R. Knox and Dr. E. L. Ghisalberti have played a
major role in the chemical work, much of which was
carried out by research students in the Department.
We are also grateful to Mr. J. Matisons for technical
assistance.

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107

Journal of the Royal Society of Western Australia, Vol. 62, Parts 1-4, 1979, p. 109-128.

Aspects of Science in Western Australia 1829-1979.

A vclume to commemorate the 150th Anniversary of the founding of Western Australia.

The prehistoric environment in Western Australia

by D. Merrilees

C/- Western Australian Museum, Francis Street, Perth, W.A. 6000

Abstract

Knowledge of various aspects of the local environment, excluding climate (see
Wyrwoll, this volume) is reviewed for an arbitrarily chosen period of 200 000 years.
Such knowledge is extensive, though patchy, and this combined with the need to
fit it into the extensively but patchily understood world scene puts comprehensive
reviewing bevond the range of one man. Consequently the review attempts merely
to indicate the scope of studies of some aspects of the environment, any one of
which might now be reviewed comprehensively with advantage. An attempt is
made to review prehistoric mammals comprehensively, and some new information is
reported.

So far as present knowledge extends, it is concluded that direct and indirect
effects of glacioeustatic changes in sea level, and changes in the mammal fauna
would provide the most noticeable contrasts with historic time. For most macro-
scopic forms of life expansions and contractions of range of familiar species have
been characteristic of the prehistoric period rather than changes in form or size.
For mammals (and probably for other less well-known groups) extinction without
replacement was also characteristic, at times and for reasons which remain obscure,
though human influence is suspected. Some changes in range appear to have been
vegetationally and hence probably climatically induced, and one or two might be
due to interspecific reaction.

Introduction

This review of the prehistoric environment,
other than climate (reviewed by Wyrwoll, this
volume), is made from the point of view of a
human observer travelling backward in time and
noting (by simple use of his senses) differences
from the environment of his temporal starting
point. For convenience, this is taken to be the
environment as the first European settlers saw
it, not as is seen at the time of writing after 150
years of drastic modification.

This hypothetical journey through time is con-
ceived from the results of technologically-
sophisticated investigation, of which there has
been a great deal, so that a complete review
might become merely an annotated list of studies
of late Quaternary phenomena, not only in
Western Australia itself, but elsewhere as they
affect this large region. To avoid this and so
produce a readable short review, I have perhaps
fallen into the trap of being altogether too
sketchy and selective. But except in the section
on prehistoric mammals, there is no attempt to
be comprehensive, merely to draw attention to
kinds of studies, any of which might be reviewed
comprehensively with advantage.

The “prehistoric period” is here taken to begin
with the first arrival of hominids in what is now
called “Western Australia”, which may well be
their first arrival in any part of Australasia.

This time cannot yet be fixed with any confi-
dence, and so it is necessary to choose a starting
point arbitrarily. I choose 200 000 years, know-
ing that this will seem far too early to many
readers and that it is in excess of the estimate
of Bowler (in Kirk and Thorne 1976) but finding
it convenient and believing it to be reasonable,
if not, indeed, too late.

A number in brackets in the text after a
locality name means that the locality is indicated
in Figure 1. Specimen numbers are those of
the Western Australian Museum collections.

Chronological framework

Many phenomena have been shown to have
similar periodicity, some no doubt because they
are causally related. But whether there were
many episodes of about the same amplitude, or
a few of large amplitude with many minor oscil-
lations, as implied in the traditional Alpine ice
age terminology, is still debatable — e.g. compare
Fink and Kukla (1977) with Grootes (1978).

Following Harmon et al. (1978b), dealing with
Bermuda, I adopt “a picture of long major
periods of low sea stand” broken by briefer high
level periods, with rapid transition from one to
the other (e.g. Broecker and van Donk 1970,
Berger et al. 1977, Yapp and Epstein 1977). This
accords well with studies as diverse as those of
Harmon et al. (1978a) on North American

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Journal of the Royal Society of Western Australia, Vol. 62, Parts 1-4, 1979.

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Journal of the Royal Society of Western Australia, Vol. 62, Parts 1-4, 1979.

speleothems, Luz (1977) on South Pacific plank-
tonic foraminifers or Woillard (1978) on Euro-
pean pollen. Shorn of the obscuring effects of
minor oscillations, it is well seen in the study of
sea floor sediments off New Zealand by Lewis
(1973).

For present purposes I adopt the following
synthesis, based mainly on the core near Perth
studied by Be and Duplessy (1976), recognising
that there is much controversy about the rela-
tive durations of periods of high and low sea
level, and about the dates of transitions (the
beginning and end dates are arbitrary) :

Sun, moon and stars no doubt would have
seemed constant throughout our prehistoric
period. Differing relative movements related to
the precession of the equinoxes might have been
noticed by a seafaring people like those of the
Pacific islands, but perhaps not by the contin-
ental people of Western Australia. But at least
one apparently celestial event might have been
spectacular, namely the tektite shower, part of
it considered by Lovering et al. (1972) to have
fallen into active seif dunes between 24 000 and
16 000 years ago near Lake Torrens in South
Australia.

0-10 x 10 3 yr B.P.
10-100 x 10 3 yr B.P.
100-130 x 10 3 yr B.P.

130-190 x 10 3 yr B.P.
190-200 x 10 3 yr B.P.

Warm, sea initially rising rapidly
to level slightly above present,
then falling.

Cool with low sea levels — minima
a little after 70 000 and 20 000 yr
B.P., the latter very marked.
Warm, high sea levels — maxima
a little higher than the present
at about 120 000 yr B.P. or a
little earlier.

Cool, low sea levels — minima
about 150 000 yr B.P.

Final phases of a warm period —
sea level falling rapidly.

Extra-terrestrial events

Many of the climatic effects impinging on pre-
historic man no doubt had extra-terrestrial
origins, some perhaps resulting from fluctuations
in solar output. Sunspot activity may be linked
with energy output and hence with terrestrial
events, as suggested by Eddy (1976). It is even
possible that more distant stars or interstellar
material have terrestrial effects, as in the con-
cept reviewed by Dennison and Mansfield (1976),
or that of Hunt (1978).

Some effects of immediate terrestrial origin
may have interacted with extraterrestrial effects,
even if not causally related. For example, there
have been several recalculations and much sup-
port (e.g. Veeh and Chappell 1970, Emiliani and
Shackleton 1974, Weertman 1976, Hays et al.
1976, Berger 1977) for concepts of this kind
associated particularly with Milankovitch (Kukla
1976). Whyte (1977) has suggested that decel-
eration of mantle spin velocity is related to
reversal of geomagnetic polarity and this in
turn to decreasing oceanic ridge activity, marine
regression and climatic “deterioration” leading
to glaciation. The climatic effects may be due,
at least partly, to changes in ionised layers sur-
rounding the Earth with changes in its magnetic
field (Cox in Bishop & Miller 1972).

Horwitz and Hudson (1977) show that the
Western Australian tektites covered the whole
State, not just the southern part, as formerly
believed. They do not suggest a date of fall,
but imply this was the same as that of the east-
ern occurrences, with the possible exception of an
apparent indochinite chip from an archaeological
excavation at Miriwun (33) in the Ord Valley
(Cleverly and Dortch 1975) which may represent
a separate fall. There is still an unexplained
discrepancy between the isotopically determined
age of formation of australites (c. 700 000 yr B.P.
or more) and stratigraphically determined ages
of fall (late Quaternary, even Holocene) accord-
ing to Chalmers et al (1976), and until this is
resolved, such occurrences as the Murchison
River (21) tektite, apparently associated with
an extinct marsupial and artifacts (Merrilees
1968c) remain puzzling. Durrani and Khan
(1971) suggest that ecological effects might have
accompanied the fall of microtektites.

Whether or not prehistoric Western Austra-
lians witnessed one or more tektite showers, pre-
sumably as a swarm of “shooting stars”, the
material results of the shower (s) seem to have
been a valued component of their environment
(Cleverly and Dortch 1975).

Presumably also they witnessed the fall of
most or some of the numerous iron and stony
meteorites known from many parts of Western
Australia, especially the Nullarbor region (com-
pare Wasson 1974), but probably not including
crater formation at Dalgaranga (20) and Wolf
Creek (32) (McCall 1977). It would be inter-
esting to know if any of these more dramatic
occurrences achieved legendary status. Met-
eorite material, unlike tektite glass, does not
appear to have been put to mundane use, but
may have had some magical importance.

Figure 1. — Western part of Australasia at minimum sea levels of prehistoric period. Inset: emergent shape of
Australasian landmass and proximity to emergent Asia during glacioeustatic minimum sea levels. Numbers
represent localities or areas mentioned in the text; 1 — Koonalda Cave; 2 — Thylacine Hole, Horseshoe Cave-
3— Madura, Murra-el-elevyn and other caves; 4— Balladonia district, Wonberna, Guralia and other rockholes-’
5— Archipelago of the Recherche; 6— Bremer Bay district, Reef Beach, Hunter River; 7— Donnelly River Bo^gy
Lake; 8— Scott River, “Warren Beach”; 9 — South-western cave area including Devil’s Lair Mammoth Cave
Skull Cave, Hamelin Bay; 10— Various lunettes; 11— Caves south of Mandurah; 12— Garden and Rottnest
Islands; 13— York, Quairading; 14— Mirrabooka, Mosman Park, Leighton Beach, Fremantle, Guildford Orchestra
Shell Cave, Melaleuca Cave, Koala Cave; 15— Gingin district including McIntyre Gully and Caladenia Cave-
16— extinct rivers; 17— Jurien Bay region including Hastings and Weelawadji caves; 18— Abrolhos Islands’
19— Greenough River, Merkanooka; 20 — Dalgaranga meteorite crater, Wilgia Mia; 21 — Murchison River sites
including Billabalong; 22— Shark Bay region; 23— Warburton region including Puntutjarpa rock shelter-

24 — Gascoyne River; 25 — Lake McLeod; 26 — North West Cape region including Monajee Cave- 27 Pilbara rock

art sites; 28 — Cossack district; 29 — Barrow Island; 30— Fitzroy River EsLuary; 31 — Windjana Gorge Tunnel

Creek; 32 — Wolf Creek meteorite crater; 33 — Ord River region including Miriwun rock shelter- 34 To<;pnh

Bonaparte Gulf; 35— Alligator River rock art sites. ’ ep

111

Journal of the Royal Society of Western Australia, Vol. 62, Parts 1-4, 1979.

Volcanic and tectonic activity

Unlike western Victoria (Gill 1972) or the
Carpentarian coast (Grimes and Doutch 1978)
for example. Western Australia does not appear
to have experienced any volcanic activity in the
prehistoric period, although of course, indirect
effects of volcanicity elsewhere may have been
felt climatically (compare Ninkovich and Donn
1976, Bray 1977).

Tectonic activity in this State appears to have
been on a much smaller scale than that in the
New Hebrides (Neef and Veeh 1977), for example.
No doubt earthquakes occurred in the prehistoric
as in the historic period, and may be recorded
in legend (Hallam 1975), but no geological traces
of them seem to have been recorded. Slower
movements persisting into prehistoric, perhaps
even into Holocene time, have been reported or
suggested by Cope (1975), Playford and Cock-
bain (in Walter 1976), Playford (in Playford
and Leech 1977), Denman and van de Graaff
(1977) indicating a need for caution in inter-
preting glacioeustatic markers round the coast
(van de Graaff 1978). As with volcanism, it is
possible that distant tectonism had effects here
(compare Roosen et al. 1976).

Coasts and islands

Glacioeustatic low sea levels probably would
produce coastal changes noticeable by our hypo-
thetical time traveller, particularly where he had
some stable and familiar marker of position, such
as the Darling Scarp provides along the south-
western coast. Since the present is a time of
glacioeustatic high sea level, with only two com-
parable levels in the last 200 000 years, the effects
of these probably would be scarcely noticeable
(cf. Tankard 1975, Marshall and Thom 1976,
Smart 1977).

The approximate coastline of the minimum
prehistoric sea level (c. 17 000 yr B.P.) is plotted
in Figure 1 on the assumption that the blanket
of sediment deposited subsequently has not
altered the once emergent landscape to any
great extent, and that glacioeustatic movements
are of about the order of magnitude reported by
Chappell and Thom (in Allen et al. 1977).

The great extension of the northern Australian
lowlands during times of low sea level is note-
worthy, and effects on prehistoric human popu-
lations and on climate would be profound. The
lesser extension of the south-western and south-
ern coasts might still have affected local people
appreciably, as in the case of Devil’s Lair (Balme
et al. 1978).

All islands (and reefs) at present within about
150 km from the Western Australian coast would
have been parts of the mainland at least twice,
more likely three times, within our arbitrary
“prehistoric” period, and twice or three times
isolated, including the present high sea level
period of isolation.

Incorporation with the mainland and isolation
of islands have been discussed, in several cases
in the context of effects on animal and plant
populations, in the specific cases of Barrow
Island (29) (Main and Yadav 1971), the shore

near Lake McLeod (25) (Denman and van de
Graaff 1977), Bernier, Dorre and Dirk Hartog
Islands (west of 22) (Kendrick 1978), the Abrol-
hos (18) Teichert 1947, Storr 1965), Garden
and Rottnest Islands (12) (Churchill 1959, Play-
ford and Leech 1977) and the Recherche Archi-
pelago (5) (Fairbridge and Serventy 1954, Main
1961). Only the most recent of these studies
have been able to take tectonic as well as glac-
ioeustatic movements into account.

Submarine canyons off the present southern
Australian coast are discussed by von der Borch
(1967). Most of the major ones seem to have
been cut long before our prehistoric period, and
presumably would have been established and
noteworthy features of low sea level coasts during
this prehistoric period. A brief review of south-
ern continental areas now submerged is given
by Playford et al. (1976).

Rivers, lakes and springs

Most of the sporadically flowing or extinct
rivers and dry lake beds often given undue
prominence in maps of Western Australia seem
to have had their genesis at least as early as
the Cretaceous, and to have ceased to carry water
by the Miocene (van de Graaff et al. 1977) except
temporarily. Thus they had assumed their pre-
sent aspect long before our prehistoric period
began, and in this long perspective, prehistoric
fluctuations probably were minor. Nevertheless,
such fluctuations might have been important to
the prehistoric people.

For example, Kendrick (1977) shows that the
Swan estuary supported an essentially marine
fauna at least as far upstream as Guildford (14)
in mid-Holocene time. The regional aridity
implied by this no doubt had far reaching human
effects, and one immediate effect would have been
on the quality and quantity of food available to
river bank dwellers. Not until some time after
4 500 B.P. was the existing hydrologic seasonality
of the river established (or re-established).

An apparent association of prehistoric man
and the large extinct marsupial Zygomaturus in
the middle reaches of the Murchison River (21),
reported by Merrilees (1968c), has been investi-
gated recently by Clarke and Lofgren, who sug-
gest (pers. comm.) that the association is real
and that it dates from a time of higher river
discharge than the present, possibly from a
declining phase of the previous major inter-
glacial, of the order of 100 000 years ago. A new
discovery of the same association has been made
in the Greenough River (19) (Dortch and Wyr-
woll, in press), with a suggested age in excess of
40 000 years. It is conceivable that the presence
of the large herbivore and the high river dis-
charge had a causal climatic connection of great
importance to the people concerned. (It is pos-
sible also that the occurrences of what appears
or is reported to be Diprotodon at Karonie, Lake
Darlot, the Great Victoria Desert and the Oak-
over River (Merrilees 1968c) date from lake-
or river- full episodes within our prehistoric
period, though there is no indication at present
of any association with artifacts.)

Journal of the Royal Society of Western Australia, Vol. 62, Parts 1-4, 1979.

A review of drainage patterns over the whole
of Western Australia, involving reconstruction
of ancient systems now ill defined, is given by
Mulcahy and Bettenay (1972), who recognize
six major divisions (see also van de Graaff et al.
1977). Their South-West Drainage Division is
described in more detail by Bettenay and Mul-
cahy (1972). If rivers in the usual sense did
exist during our prehistoric period in the now
arid or semiarid interior, they would have been
in the positions shown by these authors, but they
do not attempt to date river- (and lake-) full
periods.

According to Bowler (1976), many Western
Australian lake beds may have held water during
the prehistoric period, but it is likely to have
been saline. Many have been dry long enough
for elaborate human use, e.g. the stone arrange-
ments on Lake Moore (16) (Gould and Gould
1968).

Modern groundwater phenomena have been
extensively examined in some areas, but not
often dated. However, the subsurface structures
in the Mirrabooka area (14) north-east of Perth
(Allen 1977), for example, which determined the
sites of modern springs and soaks presumably
has changed only at the most superficial level
since the prehistoric period began, and the
springs and soaks concerned, though varying in
volume as effective precipitation varied (Wyrwoll,
this volume), might have persisted through this
period. Similarly, if the Bassendean Dune Sys-
tem is indeed earlier than “prehistoric”, as
McArthur and Bettenay (1960) suggest, then the
interdunal lakes, swamps and seasonal streams
which are very prominent on these dunes over
most of the Swan Coastal Plain might also have
persisted. But presumably the terrestrial water
table responded to glacioeustatic sea level move-
ments in ways not at present clear, and our
understanding of ground-water effects is further
clouded by large scale destruction of the natural
plant cover for agricultural and other purposes
and by the relationship of present ground water
to past aridities (Kendrick 1976).

Rocks, landforms, soils

Australia separated from the rest of Gondwana-
land, the Indian Ocean came into being, and the
basic geological structures which determine land-
forms were in existence long before our prehis-
toric period, as succinctly outlined in Geological
Survey of Western Australia (1975) and, with
special reference to the south-west, by Playford
et al. (1976). Against such a background, pre-
historic changes could only have been on a mmor
scale.

Differences from the present most easily recog-
nised by a time traveller probably would be
those associated with glacioeustatic low sea
levels. Thus Clarke and Dortch (1977) present
an early Holocene view of their Minim Cove
(Mosman Park) locality as perched on a high
limestone ridge overlooking the deep gorge of
the Swan River (14), presumably in transition
from even more extreme dissection a few thou-

sand years earlier, with a coast about 50 km
away, to its present state of low relief, broad
estuary and coast within 2 km.

Another noticeable by-product of glacioeustatic
movements, including those of prehistoric as well
as earlier Quaternary time would be the aeolian-
ites of the Tamala Limestone. A long series of
studies on similar aeolianites on Bermuda, sum-
marized by Harmon et al. (1978b) has tended to
confirm early concepts of the source material
having been produced in the warmed seas of
interglacial times, to be transported landward
in times of retreating seas, and lithified during
times of low sea level. If so, we must envisage
at least three dune generations within our pre-
historic period, and such dunes are likely to
have transformed the local landscape. Studies
of a small area near Fremantle (14) (Kendrick,
in prep.) are likely to test the local applicability
of these Bermudan concepts; these studies hinge
on the interpretation of the two late Pleistocene
radiocarbon dates mentioned by Playford et al.
(1976).

The importance of caves and cave deposits is
so great to the prehistorian (Bowdler 1975) as
to tempt him to overestimate the importance of
caves to prehistoric people. Yet these people
clearly did use caves, not only shallow rock
shelters such as Miriwun (33) (Dortch in Wright
1977), but also deep limestone caves. Perhaps
the most dramatic case is that of Koonalda
Cave (1), just outside Western Australia (Wright
1971), but Balme et al. (1978) and Clarke
et al. (in press) show that the present small
extent of Devil’s Lair (9) is misleading about
its depth when it was in prehistoric human use.
It may be that the midden locations in Mam-
moth Cave (9) (Archer et al. in press) were
deeper within the cave when they were in use
than they are now, through downstream migra-
tion of the entrance. Many other caves were
used by prehistoric groups, some apparently for
domestic purposes, others (e.g. Orchestra Shell
Cave (14), Hallam 1974) perhaps for ritual use.

Caves are very numerous in Western Australia
now, and presumably were throughout the pre-
historic period. Descriptions are far too num-
erous to cite here, but lists with brief individual
notes and full references are given for various
regions, notably the Augusta to Calgardup Brook
district (9) by Bridge (1973) and Bridge and
Shoosmith (1975), for the Yallingup area by
Bridge (1972), for the Moore River to Jurien
Bay (17) by Shoosmith (1973) and Bridge (1973)
and for Cape Range (26) by Shoosmith (1977).
Nullarbor caves are listed by Dunkley and Wigley
(1967), with some emendations by Lowry and
Jennings (1974). There appears to be no review
or list of Kimberley caves, but Jennings and
Sweeting (1963, 1966) and Lowry (1967) describe
large caves and briefly mention others.

Karst phenomena other than caves are also
common in Western Australia, for example the
depressions now occupied by salt lakes on Rott-
nest Island (12) (Playford and Leech 1977) and
circular depressions on reef flats off the Abrolhos
Islands (18) (Playford et al. 1976), interpreted
as glacioeustatic low sea level products.

113

82710 ( 8 )

Journal of the Royal Society of Western Australia, Vol. 62, Parts 1-4, 1979.

Dominant modern soils, usually correlated with
landforms, have been mapped for the whole con-
tinent in an “Atlas of Australian Soils” pub-
lished by CSIRO in association with Melbourne
University, with a series of explanatory notes for
the various regions (e.g. Northcote et al. 1967
for the south-west). Each of the series has an
extensive reference list. Presumably the soils
now present in most places have taken a long
time to develop, and many may be prehistoric in
origin, but in general there is little information
available on the ages of soils. However, ancient
soil structures or soil remnants feature prom-
inently in the Western Australian landscape, for
example the pedogenic carbonate, distinguished
(as “kankar”) from valley fill “calcrete” by
Sanders (1974), which is very widespread in the
dry interior. This may well be much older than
our “prehistoric period”.

Buried soils occur in the Tamala Limestone,
for example those at Hamelin Bay (9) described
by Fairbridge and Teichert (1953). They are
often well defined, sometimes well lithified, and
may be prominent features of cliffs, or form the
roofs of caves, but even so represent short-term
highly localized episodes of soil formation in
many cases, often in interdunal swales (Clarke
et al., in press). There are additionally very
densely lithified bands often with solution pipes
descending from them, sharply demarcated from
overlying non-calcareous sands. Whether these
lithified bands (“cap rock” or “capstone” in
local lay usage) are horizons in a soil profile
including the overlying sand, or are completely
separate (as in the site described by Clarke and
Dortch 1977) is not always clear. Solution of
local problems such as this would seem to be
amenable to closer study, and there is a prospect
of working out a dune and soil sequence, as has
been done in Bermuda (Harmon et al. 1978b),
which with marine intercalations will permit-
stratigraphic subdivision of the Tamala Lime-
stone. In the meantime, it is not clear which
parts, if any, are “prehistoric” in our sense.

Relative soil and landform chronologies have
been put forward for some regions (e.g. for York-
Quairading (13) by Mulcahy and Hingston
1961), and an attempt made to establish an
absolute chronology for the Swan Coastal Plain
by McArthur and Bettenay (1960). Though
subsequent refinements in the world Quaternary
chronology on which this is based may suggest
some revision, it is likely that many of these
Swan Coastal Plain soil systems can still be
regarded as “prehistoric” in origin.

South-western soils are reviewed by Mulcahy
(1973).

Deserts and dunes

With his convenient though European-oriented
definition of “desert” Beard (1969) points out
that about two fifths of Western Australia is
involved and suggests natural divisions. These
would seem to have been in existence long before
our prehistoric period began, but nevertheless
in their extreme form may be Quatenary phen-
omena, by analogy with the eastern Australian
occurrences discussed by Jessup and Norris
(1971), Bowler (1976) and others.

If so, the perspective is shorter than for rivers
and lakes, and one might expect prehistoric
fluctuations in deserts to have had profound
effects on prehistoric human populations in-
volved with them. Even now, most desert and
desert-fringe dwellers not only in Australia but
elsewhere seem to be more at the mercy of this
than of most environments, and yet to influence
it more profoundly by their mere existence,
usually to accentuate if not actually produce
desertification (Hare 1977).

By analogy with other parts of Australia (e.g.
western New South Wales, Wasson 1976), it
would seem that some desert dunes and even
sand sheets of modest size might still have been
mobile almost to the end of our prehistoric
period, and Jennings (1975) and Bowler (1976)
demonstrate dune building on a large scale in
late Pleistocene time.

One special kind of desert or semidesert land-
form, the lakeside lunette, seems to have had
considerable importance for prehistoric people.
Investigations of lunettes in the Murray-lower
Darling River region are described by Bowler
(1970, in Mulvaney and Golson 1971, 1976), Mer-
rilees (1973a), Marshall (1973), Allen (1974)
and others. They have received less attention
so far in Western Australia, and may have been
less important prehistorically, but some aspects
are discussed by Bettenay (1962) and Bowler
(1976) (10).

The Western Australian deserts were not
empty of people for the whole prehistoric period.
Gould (1968, 1977) demonstrates occupation

dating back about 10 000 yr., continuing into the
historic period. But it is possible that our deserts
were only habitable during interglacials, if
Gould’s stress on the difficulties they now present
is in proper perspective.

Air and sea

There is reason to think that the present
chemical composition of the Earth’s sea and air
envelopes is the current stage in a long sequence
of evolutionary changes (e.g. Fairbridge in Chil-
ingar et al. 1967), but that there has been no
catastrophic change in late Cainozoic time.
McLean (1978) suggests that land plants have
been the major regulators of carbon dioxide and
oxygen in the air throughout the Cainozoic, and
that the effect of major marine transgressions
would be to increase carbon dioxide and concom-
itantly decrease oxygen content of the air by
replacing large areas of terrestrial with marine
floras. Possibly reinforcing this effect is another
described by Berger et al. (1977), a release into
the air of carbon dioxide previously trapped in
deep ocean water under a “lid” of glacial melt-
water at the time when vertical mixing became
more effective as deglaciation (and hence marine
transgression) proceeded. In view of the import-
role of the relatively small quantities of carbon
dioxide now present in the air (Bray 1977), it
is very likely that prehistoric people would have
been affected indirectly by this change two or
three times during the prehistoric period.

Fluctuations in the rate of removal of calcium
from sea water by plants and animals presum-
ably occur in response to the relatively small

114

Journal of the Royal Society of Western Australia, Vol. 62, Parts 1-4, 1979.

temperature increases (Be and Duplessy 1976,
Shackleton and Opdyke 1977) of interglacial
times, as implied or stated in various adaptations
of the Sayles model of dune limestone formation
(e.g. Sayles 1931). But perhaps they are oal-
anced by inverse fluctuations in delivery of cal-
cium to sea water by river discharge. On the
other hand, if chlorine and bromine are con-
tinually discharged into sea, air and sediments
(with the sea accounting for a large proportion
of this “surface reservoir”) by volcanic processes
(Schilling et al. 1978), then the quantities pre-
sent will have altered over the prehistoric period.
But the rate of alteration presumably would be
perceptible only by sophisticated means.

Fluctuations in particulate matter, both in sea
and air, might be directly observable. Thus the
air of glacial time might have been perceptibly
dustier than in historic time (compare Kolia
and Biscaye 1977), and smokier in late than
in early prehistoric time (Hallam in Wright
1977). A significant and environmentally
informative component of dust, not yet studied
in Western Australia, is phytoliths (Baker
1959a, b, Smith et al. 1970, Rovner 1971). There
is still controversy about the sediment load of
rivers of glacial compared with interglacial time
(Flint 1971, Verstappen 1974), but there might
well have been differences, affecting among other
easily observable things, the colour as well as the
turbidity of the sea, and the distribution of
molluscs and other organisms important in pre-
historic human diet.

Physical processes in the air of the kind usually
summed up as climatic fluctuations are discussed
by Wyrwoll (this volume). Similar processes in
the more viscous material of the sea might be
expected to be slower, but to have no less pro-
found effects, not least through their influence
on atmospheric processes (e.g. Berger et al. 1977).
Wyrwoll (this volume) suggests that surface
currents have been significant determinants of
climate in Western Australia.

Animals

Mammals

The most recent review of Quaternary mam-
mals in Western Australia is that of Merrilees
(1968c), which was not intended to be confined
to the “prehistoric period” as defined here, but
which in fact largely was so confined. By adding
the following notes of new discoveries and
studies, an up to date review is obtained.

Rock 'paintings and carvings, and traditions . —
Representations of thylacines have been noted
by Wright (27) (1968, 1972), Brandi (35) (1972,
1973) and Lewis (1977), and of devils by McCar-
thy (1976) and Calaby and Lewis (1977), though
many of these are outside Western Australia.
These and earlier records of extinct forms, coup-
led with renewed interest in and appreciation of
Aboriginal art (Edwards and Ucko 1973, Ucko
1977. Clegg 1978), might be expected to lead to
systematic search for representations of totally
or locally extinct taxa, comparable with those
of other countries (e.g. Bruemmer 1974, Powers
and Stringer 1975).

An oral tradition which might relate to the
thylacine (Kolig 1973) exemplifies another
avenue of investigation of the prehistoric fauna
and the use of Sthenurus and Zygomaturus teeth
in charms (Akerman 1973) still another. The
latter occurrence does not necessarily imply co-
existence of man with Sthenurus and Zygoma-
turus, as interest in and use of fossils by Aborig-
ines seems to have been widespread (White-
house 1948, Gill 1957), as by other prehistoric
people (e.g. Oakley in Brothwell and Higgs 1969,
Kraft and Thomas 1976). Even now, Aboriginal
people maintain this interest (M. Lofgren, pers.
comm.).

Fossil deposits in the South-West . — Knowledge
of these has been extended greatly since the
review by Merrilees (1968c), and a new interpre-
tation of the Mammoth Cave (9) deposit put
forward by Archer et. al. (in press).

Mammoth Cave now has a walk-in entrance,
and probably has had for most or all of the
period under study, though perhaps not always
in its present position. As with most caves in the
region, there is a surface scatter of bones, pre-
dominantly of mammals with several local con-
centrations. But there are (or were) also two
patches rich in bone buried in partly lithified
deposits capped by thick flowstones. These were
dug out in the early years of the present century
leaving only small remnants, and some of the
bone was described from a taxonomic viewpoint,
with little attention to provenance, in a series
of papers by L. Glauert (see Glauert 1926, under
“Pleistocene”). Glauert’s findings have been
quoted frequently (e.g. by Tedford 1967), but
require extensive revision. Some was made by
Merrilees (1968c) and some further revision is
incorporated in Table 1. Merrilees and Porter
(in press) illustrate and give diagnostic char-
acters of some Mammoth Cave fossil material,
and tentative reconstructions of some taxa are
included in Figure 2.

The two richly fossiliferous patches in Mam-
moth Cave have been distinguished as the “Le
Souef” and “Glauert” deposits respectively by
Archer et. al. (in press), but material from them
was not kept separate by the excavator, and
little can be distinguished now. Though sep-
arated topographically by about 10 m, the two
deposits appears to be strategically continu-
ous. Several layers can be distinguished in the
remnant of the “Glauert deposit” and at least
two in the “Le Souef deposit”, but material from
these layers was not labelled separately, and only
some specimens can be distinguished with con-
fidence as from one or other layer. The lowest
layer, probably the one containing all or most
of the extinct taxa, has an age beyond the pre-
sent practical range of radiocarbon dating
(>37 000 radiocarbon yr B.P.; Merrilees 1968c).

As a result of studies of a variety of marks on
the bones, Archer et. al. (in press) conclude that
a notch on a fragment of Sthenurus tibia is
man made, and conclude also that extensive
breakages of large limb bones are artificial.
Some of the bone is charred and two fragments
possibly represent human skull, but no stone
tools have been detected so far. From this

155

Journal of the Royal Society of Western Australia, Vol. 62, Parts 1-4, 1979.

Table 1

Revised list of vertebrate remains from Pleistocene
deposits in Mammoth Cave, i.e. from “Glauert” and
“Le Souef” deposits (Archer et al. in press), ignoring
undated material from elsewhere in the cave.

Monotremes

Tachyglossus aculeatus e.g. 61.7.14

Zaglossus hacketti (= Z. bruijini?) .... e.g. 60.10.1
Echidna of intermediate size e.g. 61.6.6

Carnivorous marsupials of medium or small size

Dasyurus geoffroyi

.... e.g.

66.9.10

Phascogale tapoatafa

.... e.g.

66.9.7

Antechinus flavipes

.... e.g.

68.6.285

Sminthopsis murina

.... e.g.

68.6.286

Sarcophilus harrisii

.... e.g.

66.9.14

Thylacinus cynocephalus

.... e.g.

61.2.25

Bandicoots

Isoodon obesulus

.... e.g.

66.2.195

Perameles (P. bougainville? )

.... e.g.

66.7.12

Possums, koala, wombat, etc.

Trichosurus vulpecula ....
Pseudocheirus peregrinus
Thylacoleo (T. carnifex?)
Phascolarctos (probably P.
Vombatus hacketti

e.g. 70.3.27

e.g. 69.3.767

e.g. 64.10.25

cinereus) .... e.g. 64.2.26
e.g. 65.1.21

Macropods

Potorous tridactylus

Onychogalea (possibly O. unguifera)

Macropus eugenii

Macropus irma

*Macropus fuliginosus

Setonix bracliyurus

“Mammoth Cave wallaby" — an unidenti-
fied, probably undescribed species about

e.g. 66.3.65
e.g. 69.3.831
e.g. 69.3.637
e.g. 66.9.36
e.g. 69.2.241
e.g. 69.3.219

the size of Wallabia bicolor or Macropus
agilis, represented by juvenile dentaries,
e.g. 66.8.17. Adult maxillary fragments
(e.g. 66.8.18) may or may not represent
the same species

Sthenurus brownei

e.g.

63.2.198

Sthenurus occidentals ..

.... e.g.

62.8.31

Protemnodon brehus

e.g.

64.2.18

( ?Protemnodon larger than P.

brehus — e.g.

65.4.153)

Diprotodontid

Zygomaturus trilobus

e.g.

61.8.3

Eutherians

Bat

e.g.

71.3.64

(?Pseudomys albocinereus

e.g.

76.2.92)

Pseudomys shortridgei ....

e.g.

76.2.94

Rattus fuscipes

e.g.

72.4.1

Birds

Small species

e.g.

67.7.1

Eagle-like species

e.g.

61.6.12

Emu-like species

e.g.

65.4.152

Reptiles

Wonambi naracoortensis

e.g.

68.4.30

Lizard

e.g.

76.2.86

Amphibians

( ?Frog

e.g.

76.2.96)

* A second species similar in size to M. fuliginosus is
reported by Merrilees (1968c). No detailed statistical
analysis has been made, but it now seems likely that
all the large macropodine cranial material could be
included under M. fuliginosus. The possibility that
M. giganteus might be present has not been investi-
gated.

evidence and from the position of the material
on top of large blocks of limestone fallen from
the roof of the cave, Archer et al. suggests that
man was the main bone accumulator in this case.

Bone in the Devil’s Lair deposit also appears
to be the leavings of predators, mainly man in
the upper layers and owls in the lower, with an
intermediate zone of overlap. Analysis of this
bone, representing uninterrupted but slow
accumulation over the period approximately
from 35 000 to 5 000 yr B.P., has been made by
Balme et al. (1978). They report fluctuations
in the relative proportions of 23 mammal species,
and sporadic presence of another 11. For pre-
sent purposes, the most noteworthy of these
fluctuations appears to be marked decline in
latest Pleistocene and early Holocene time of
a group designated as “non forest” mammals
( Perameles , Bettongia lesueur, Petrogale , Pseu-
domys albocinereus , P. shortridgei and Notomys )
with concomitant increase in Potorous and
Setonix.

Remains of extinct mammals (e.g. Sthenurus
brownei) represented at Mammoth Cave, and of
others like Phascolarctos , which vanished from
the region before historic time, are present in
the lower layers in Devil’s Lair (9), but it seems
likely that these are secondarily derived from
another deposit of unknown age, beyond the
usual range of radiocarbon dating (Balme 1978).

Porter (1979) describes a deposit in Skull
Cave (9), spanning all or most of the Holocene,
with bones apparently representing animals
which fell into what was essentially a large pit,
could not escape, and so died. But there may
have been a contribution from owls. All the
mammal taxa present were still extant in his-
toric time, though not all were present in the
district. There are undated deposits elsewhere
in Skull Cave which on field evidence seem to be
older than that described by Porter (1979), and
Zygomaturus is represented in these (R. Howlett,
pers. comm.) ; they are still under investigation.

Small cavities in cliffs at Deepdene, which
seem to have served as owl roosts, are described
by Archer and Baynes (1973). Mammal remains
in these represent the small mammal fauna of
the region only a few hundred years before the
early historic period.

Occurrences of Petrogale in south-western cave
deposits have been described by Merrilees (1979),
and new discoveries of extinct taxa have been
made in The Labyrinth (Merrilees 1969), at
Hamelin Bay ( Vombatus , specimen 77.8.102) and
in Crystal Cave ( Protemnodon , probably P. bre-
hus, 78.2.4 and Sthenurus sp., 78.2.3.) (All 9).

Coastal dune sites along the south coast have
been systematically searched for mammal re-
mains in connection with a determined attempt
by the Department of Fisheries and Wildlife to
locate living colonies of Potorous tridactylus.
Large numbers of Setonix specimens were col-
lected, with smaller numbers of other extant
taxa, not yet reported in detail. Butler (1969)
reports mammals (including Sarcophilus ) from
such a dune (8), undated, and Butler and Mer-
rilees (1970) list a fauna including Potorous
platyops and Antechinus apicalis from another
dune (6), considered to represent owl prey and
to date from about 1 000 yr B.P. Another owl
pellet deposit in a small cliff cavity at the mouth
of the Donnelly River (7), briefly mentioned by

Journal of the Royal Society of Western Australia, Vol. 62, Parts 1-4, 1979.

At least three species
\ of native rats and mice

Thylacinus cynocephalus

Wonambi naracoortensis

Zaglossus hackctti Larger Tachyglossus

Figure 2.— Some prehistoric inhabitants of the extreme South-West, now extinct there. All to same scale.

117

Journal of the Royal Society of Western Australia, Vol. 62, Parts 1-4, 1979.

Archer and Baynes (1973 p.88), has not been
examined in detail. Merrilees (1970b) reports the
finding of Zygomaturus in dune limestone near
Bremer Bay (6); charcoal collected with this
specimen has been dated subsequently as greater
than 32 000 yr B.P. (GaK - 2542, Kigoshi et al.
1973).

Further north, the wheat belt remains virtually
blank palaeontologically, and the small caves of
the coastal region between Perth and Bunbury
have been relatively unproductive, despite sys-
tematic search by speleologists. However, Bet-
tongia lesueur (76.4.9) has been found near Man-
durah (11) presumably of prehistoric age.

In the metropolitan region, there have been
new finds of Protemnodon brehus (76.3.2) near
Leighton Beach (12), and Sthenurus occidentalis
(76.6.19) at Melaleuca Cave (15), Yanchep
National Park, both of unknown age. Archer
(1974) describes a late Holocene prehistoric
fauna (including Onychogalea lunata) from
Orchestra Shell Cave (15), Wanneroo and
Archer (1973) records what appears to be
an older deposit, with remains of Sthenurus
brownei , Potorous platyops, Phascolarctos, Vom -
batus and Perameles from Koala Cave (15),
Yanchep. This may or may not be prehistoric.
Amendments to the account by Douglas et al.
(1966) of a deposit at Wanneroo (15) include the
recognition in it of at least one artifact (Merri-
lees 1973b), and of Onychogalea (probably O.
lunata — 65.10.164, 188, 190) among specimens
previously misidentified as Petrogale.

Further north, excavations in Hastings Cave
(17), not yet reported, have shown substantial
changes in the mammal fauna during the Holo-
cene, and have confirmed the presence of Pro-
temnodon brehus (72.12.173) (A. Baynes, pers.
comm.). Excavations in Caladenia (15) and
Weelawadji Caves (17) have not yet been re-
ported, and new finds have been made in the
Gingin district (15) where Tyrer (1971) reports
a Protemnodon molar fragment of indeterminate
species (69.9.10 is a cast of Tyrer’s specimen),
and Roe (1971) records a late prehistoric or early
historic fauna from a small excavation.

Repeated visits to the Zygomaturus locality in
McIntyre Gully (15), Gingin (Merrilees 1968c
p.13) by R. Roe, F. Dodds and others have pro-
duced a series of new finds, including further
Zygomaturus specimens, one reported by Merri-
lees (1970b). Specimens from McIntyre Gully
not so far reported include a dentary fragment
(78.7.2) representing an apparently undescribed
species of Sthenurus larger than S. occidentalis.
Post-cranial fragments tentatively ascribed to
Sthenurus , Protemnodon, Macropus cf. M. fulig-
inosus, and a medium sized macropodine have
been recovered, as well as a small number of
specimens representing Macropus irma, Betton-
gia (probably B. penicillata) , Setonix, Isoodon
and Dasyurus. The specimens concerned are
78.7.2-25, 75.12.2 and 69.4.48-52.

The geological character of the McIntyre
Gully deposit remains obscure, and material of
modern and Cretaceous age is mixed with pre-
sumed Pleistocene material, not always readily
separated by appearance. There are shallow

incisions on at least one fragment of a small
limb bone (69.4.50), but no certainty that tnese
are artificial and no other suggestion of human
presence.

A Zygomaturus mandible (77.9.10), with stone
artifacts from the same bed (Dortch and
Wyrwoll, in press) was discovered in the bank
of the Greenough River (19), in 1977, and a
collection of fragmented bone, representing a
variety of mammals (including Leporillus api-
calis and Onychogalea lunata) as well as birds
and other taxa was made at Merkanooka (19).
The latter collection 65.11.22-28, 66.1.6-11,

68.1.30-48 included artifacts and charred bone,
but whether of prehistoric or early historic age
is not certain.

North-West Division. — Association of artifacts
with Zygomaturus remains from the Murchison
River (21) reported by Merrilees (1968c), has
been confirmed (J. D. Clarke and M. Lofgren,
(pers. comm.). Burbidge and George (1978)
quote finds by A. Baynes and D. Merrilees of
“sub-fossil”, presumably prehistoric, mammal
remains, including Leporillus conditor on Dirk
Hartog Island (west of 22) and A. Baynes is
studying material collected by him in other parts
of the Shark Bay region.

Thylacinus and other taxa are reported from
cave deposits on North West Cape (26), at least
one of which (in Monajee Cave), has an archaeo-
logical component (Kendrick and Porter 1974).

Kimberley Division. — Archaeological deposits
of Holocene and very late Pleistocene age in the
Ord River region (33) are described by Dortch
(in Wright 1977). Some of these contain mammal
remains, not yet analysed in detail. Archer
(1975) lists Thylacinus and other mammals from
the Tunnel Creek area (31) which might be
“prehistoric”, if indeed not “historic” in time.
Small numbers of specimens of extant taxa from
scattered localities are present in the Western
Australian Museum collection, and a Protemno-
don specimen in the Queensland Museum col-
lection is reported to come from the old “Rose-
wood” station, later part of Argyle Downs (33);
these are not dated.

Eastern Division. — This vast area is repre-
sented only by very scattered specimens of
extant taxa, by the few poorly preserved speci-
mens of Diprotodon mentioned by Merrilees
(1968c) , by a Holocene archaeological sequence at
Puntutjarpa (23) containing highly fragmented
bones of extant mammal taxa (Archer in Gould
1977), and another at Wilgie Mia (20). The
Wilgie Mia collection (73.1.154-253, including
Chaeropus, Leporillus apicalis and L. conditor) is
under study by A Baynes.

Eucla Division. — Thylacines are known to have
persisted into late Holocene time in the Nullarbor
region (2, 3) (Lowry and Merrilees 1969, Merri-
lees 1970a, Lowry 1972), after the arrival there
of dingoes (Archer 1975, Milham and Thompson
1976). (Dingoes also arrived in the Cape Leeu-
win-Cape Naturaliste region before thylacines
disappeared from it (Merrilees 1979)).

An excavation in Madura Cave (3) with an
apparent association of Protemnodon, two species
of Sthenurus and Phascolarctos with an otherwise

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modern-seeming fauna and artifacts has been
reported by Milham and Thompson (1976), but
at least the Protemnodon and by implication the
other archaic taxa have been shown to be second-
arily derived. Other faunistic results of a joint
University of Sydney-Australian Institute of
Aboriginal Studies Nullarbor archaeological sur-
vey (which included the Madura Cave excava-
tion) have not been published. However, Archer
(1972) independently has listed mammal re-
mains from an excavation in one of the caves
examined in this survey, Horseshoe Cave (2)
with radiocarbon dates showing it extended back
into the Pleistocene.

Pleistocene dates (all on bone) are listed by
Lundelius and Turnbull (1973) with an ongoing
series of taxonomic studies (e.g. Lundelius and
Turnbull 1975) for still other excavations in
Madura Cave.

The Western Australian Museum collection
contains an abundance of material from Nullar-
bor caves, not yet studied in detail or reported
even at the simple “interesting occurrences’’ level
like that of Merrilees (1968d) for Chaeropus.

Lower vertebrates

Peculiarities in the modern geographical dis-
tributions of many bird taxa have led to infer-
ences about their prehistoric status. For
example, Mayr (1944) concluded that certain
species like the nankeen kestrel (Falco cench-
roides ) invaded Australia by way of Timor dur-
ing some not very remote time of glacioeustatic
low sea level, and that because most such species
were grassland birds, climatic conditions must
then have been much drier than the present.
This last logical consequence of Mayr’s thesis
flatly opposed a strong current of opinion which
then equated low latitude “pluvials’’ with high
latitude glacials, a current now largely reversed.
Fairbridge (1976) outlines the interesting history
of this reversal of opinion, and Van Devender
(1977) and Brakenridge (1978) present alterna-
tive views on . . the American Southwest . . .
one of the last areas in the world considered by
many to have experienced true glacio-pluvial
climates,” in the latter’s words.

Geologically recent arid periods have been
postulated as isolating mechanisms, leading to
speciation within the continent, by Serventy
(1951) for wrens (and Main et al. 1958 for frogs),
but pluvials also have been considered to have
this function (e.g. Ford 1974). Such postulates
usually have been linked with contemporary
opinion about the timing of climatic changes,
and there has been little study of available fossil
bird material which, if reliably dated, might yield
direct insight into prehistoric fluctuations in bird
distributions.

One fossil occurrence which may or may not
be prehistoric is that of a very large, thick-shelled
egg found by V. Roberts in a blown-out coastal
sand dune south of the Scott River (8), and
kindly loaned by the finder to the Western Aus-
tralian Museum for study and display; it is men-
tioned briefly by Rich and van Tets (1976). It
appears to be an Aepyornis egg from Madagascar.
Associated material of apparently very mixed

origin, typical of blown-out dunes, is described
by Butler (1969). This egg might have been a
collector’s item washed overboard from a passing
or wrecked ship, but it is robust, was intact when
found and could quite conceivably have floated
from Madagascar, as suggested by Lodge (1976),
a reasonable suggestion in the light of studies of
drifting objects reported by Shannon et al. (1973,
especially Figure 4). Perhaps its main relevance
here is as an example of the arrival of bizarre
material from an unknown external environment
in prehistoric time, comparable with the tektite
shower mentioned above.

Although prehistoric reptilian material is re-
corded for many Western Australian excavations
(e.g. Puntatjarpa (23), Archer in Gould 1977)
and is sometimes abundant (e.g. Devil’s Lair (9),
Balme et al. 1978), little study has been made of
it. However, the very large boid snake Wonambi
naracoortensis, initially described from south-
eastern South Australia (Smith 1976), is also
known from two Western Australian sites, Mam-
moth Cave (9) and Koala Cave (15), (M. J.
Smith, pers. comm.). A tooth from the Gas-
coyne River district (24) mentioned by Merrilees
(1968c p. 15) now appears referable to the extinct
crocodilian Pallimnarchus pollens, and like the
possibly crocodilian vertebra from the Ederga
River (19) still further south, may or may not
be prehistoric. Similarly undated crocodilian
and chelonian occurrences in the west Kimberley
region (31) are described by Gorter and Nicoll
(1978).

Study of frog remains from the extreme south-
west (Balme et al. 1978, Porter 1978) is in a very
early stage, and although fish remains have been
reported from some excavations (e.g. Miriwun,
33, in the Ord River region, Dortch in Wright
1977) and some study has been made of them,
little information is available.

Invertebrates

In abundance, variety of species and indirect
effects on people, prehistoric invertebrates no
doubt merit far more extensive study than all
vertebrates combined, as in the historic period.
Yet for lack of interest perhaps as much as lack
of durable remains, there has been very little
study of any group other than molluscs. As in
the case of birds, some writers (e.g. Richards
1971, Gray 1974) have inferred prehistoric or
earlier status from present geographical distri-
butions, but have not had the backing of direct
study of fossils.

Late Pliocene and Quaternary molluscs, both
marine and otherwise, have been reported by
Kendrick (I960, 1973, 1976, 1977, 1978 and in
Playford and Leech 1977), and he has work in
hand on a sequence of marine faunas from the
Perth Basin, some of prehistoric relevance (pers.
comm.). In general, the marine molluscs (to-
gether with corals and some other taxa) indicate
migrations or extinctions in response to tem-
perature changes, the non-marine molluscs in
response to aridity, but not all the observed
changes can be related to our “prehistoric”
period. Extinction of Bothriembryon douglasi
and B. ridei and the appearance of camaenid

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Journal of the Royal Society of Western Australia, Vol. 62, Parts 1-4, 1979.

snails in the Shark Bay region, however, is attri-
buted to a change from more to less humid con-
ditions perhaps of the order of 80 000 yr B.P.
(Kendrick 1978). Occurrence of marine molluscs
near Guildford (14) has been taken to indicate
a mid-Holocene period of aridity near Perth
(Kendrick 1977). Fairbridge (1950) discusses
the origin of coral reefs, some prehistoric.

Plants

As noted above for rivers and lakes, so for
vegetation the most profound transformation of
the Western Australian scene from one of tropi-
cal luxuriance to its familiar sparseness took
place in the early or middle Tertiary, long before
our “prehistoric” period began (Churchill 1973,
Wilde and Backhouse 1977, Kemp 1978). But
vegetational fluctuations in the prehistoric
period, though within this framework of overall
sparseness, may still have had far-reaching
effects on prehistoric people.

Unfortunately in Western Australia there have
been no studies of long pollen sequences com-
parable with those which have been environmen-
tally so informative in eastern Australia (e.g.
Dodson 1977, Colhoun 1978, Kershaw 1978). But
shorter sequences have been reported by Chur-
chill (1968) for the extreme south-west (7, 8, 9)
and Martin (1973) for the Nullarbor (3). Chur-
chill reports fluctuations with time in the distri-
bution of common eucalypt species thought to be
related to fluctuations in rainfall. Martin (and
Martin and Peterson in Pittock et al. 1978) re-
lates changes in relative abundance particularly
of chenopods and eucalypts to changes in sea
level.

Still, briefer glimpses of the vegetational past
are given by Churchill (1959, 1960 and in Hallam
1974) ; Storr et al. 1959 and Playford and Leech
(1977) for a region round Perth, including what
is now Rottnest Island (12). Jennings (1975;
reports the occurrence in mid-Holocene time of
mangroves much larger than any now known, in
the Fitzroy estuary (30), apparently related to
rainfall changes. Various sites on the south
coast, each with Holocene dates, are described by
Balme (in Dortch 1975), Bermingham et al.
(1971) and Butler and Merrilees (1971). Another
south coast site (8), not hitherto reported, is
“Warren Beach” (an informal name) nearly
west of Northcliffe, at which tree stumps, appar-
ently in growth position, can be seen submerged
for some hundreds of metres out to sea. An
extensive peat bed, possibly the substrate in
which the trees grew, seems also to be present.
Wood from one of the stumps has been dated at
8 340 ± 135 yr B.P. (SUA-343, R. Gillespie, pers.
comm.). Since sea level at this time would have
been of the order of 20 m lower that at present
and the sea floor shelves gently, the trees con-
cerned might have been as far away from the
sea as their present day counterparts are, and
there may be no climatic significance in the
occurrence. However, detailed studies have not
yet been made, nor the possibility of tectonic
movement examined (compare Cope 1975).

Parsons (1970) and Nelson (1975) studied dis-
junct plant distributions in the Nullarbor region
and the latter concludes that some represent dis-
ruption (presumably during the last major
glacioeustatic low sea level) of once continuous
distributions. Ingram (1969) reports vegetation
like the present in this region (2) in late prehis-
toric time.

Algal stromatolites, permitted to flourish in
the Shark Bay region (22) by an unusual set of
circumstances which developed in late prehis-
toric time, are discussed by Playford and Cock-
bain (in Walter 1976), and Churchill and Sar-
jeant (1962) and Harland and Sarjeant (1970)
discuss another unusual aquatic plant occur-
rence, acritarchs in the undoubtedly non-marine
environment of a deposit at Boggy Lake (7), on
the south coast.

In addition to these more or less direct studies
of plant remains, inferences have been made
about plants indirectly from studies of animals,
e.g. those of Mayr (1944), Balme et al. (1978)
and Merrilees (1978), previously mentioned.

Apart from the novelties shown by the aquatic
plants, all of these cases represent fluctuations
in floral composition or migrations of communi-
ties on a minor scale compared with the major
Tertiary transformation mentioned above.

The impact of man-made fires on vegetation
in prehistoric time is reviewed by Hallam (1975)
and suggested to have been more profound than
generally appreciated.

People

A vital component of a prehistoric person’s
environment would have been his fellow men.
The following review of this component has been
made by M. Lofgren: —

“Just how long ago hominids first arrived in
what is now called Western Australia is as yet
largely a matter for speculation. Already,
limited research completed in this vast and
diverse area has established radiometrically
documented occupational dates (9) (Dortch, in
press) comparable to the oldest dates reported
elsewhere in the continent (Bowler in Kirk and
Thorne 1976). Beyond this generally accepted
age of c.40 000 B.P. (Jones 1977) are provocative
sites and associated materials currently undat-
able by existing techniques excepting geomorpho-
logical inference. For example, sites along the
middle reaches of the Murchison River (21)
(Merrilees 1968c) may well be related to the
declining phase of the last interglacial, in the
order of 100 000 B.P. (Clarke and Lofgren, in
preparation). Such an enormous occupational
time span potentially involves a far more com-
plex human prehistory of Australia than could
have been imagined even a few years ago.

“One aspect of this complexity centres upon
the identity of those first migrants to the new
Australian world. Indeed, the timing of that
first arrival is significant in understanding vari-
ous aspects of human evolution both outside and
within Australia (Howells 1973), eventually lead-
ing to those people now called Australian Abori-
gines. While the irreducible founder population

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Journal of the Royal Society of Western Australia, Vol. 62, Parts 1-4, 1979.

might have been a young pregnant female
accidentally landing on the coast of the new
land (Calaby in Kirk and Thorne 1976), a far
more likely scenario would envision the purpose-
ful movement of coastally-adapted family groups
(Bowdler in Allen et al. 1977). Though the fail-
ure rate of such movements must have been
high, separate landing — some certainly on the
coast of Western Australia — over centuries and
indeed millenia (for possible routes, see Birdsell
in Allen et al. 1977) eventually led to the human
colonization of the continent.

“Clues as to the biological origins and subse-
quent evolutionary course of those first migrants
may be derived from a variety of sources (e.g.
Howells 1973, Kirk and Thorne 1976). Direct
evidence from studies of prehistoric human skele-
tal material, both Pleistocene and Holocene, are
vital in this regard. In the past, virtually all
these studies were restricted to material derived
from a limited area of south-eastern Australia.
Now, a recent study (Margetts and Freedman
1977) establishes a vital Western Australian
baseline against which other late prehistoric
Australian populations may be compared, as well
as providing a measure to help assess future
Pleistocene discoveries.

“While debate as to a homogeneous or hetero-
geneous origin for Australian Aborigines is
hardly new, recent studies (Thorne in Kirk and
Thorne 1976; in Allen et al. 1977; Thorne and
Wilson 1977) appear to demonstrate a much
wider morphometric range in the past. Whether
this is to be interpreted as representing separate
though related populations (i.e gracile versus
robust), or simply extremes of a more highly
variable population range remains unclear. How-
ever, new material from Western Australia (28 —
Freedman and Lofgren, in press) confirms that
this problem in morphology is not restricted to
south-east Australia. Future discoveries and
further research are necessary before the situa-
tion can be more clearly understood.

“Although their identity remains in doubt,
traces exist of their habitation of the new land
in the form of stone tools offering some insight
into human adaptions to the various Australian
environments through time. Yet, in isolation
these stone tools are of limited value in under-
standing the past (e.g. caution expressed in
various papers in Wright 1977). These tools
were secondary adjuncts to a material culture
of which little has been preserved. Indeed, the
lithic technology itself is based on ecological and
cultural grounds different from those of other
areas of the world (Hutterer in Allen et al. 1977).
Only through taking the stone technology in
conjunction with all other available data (food
refuse, pollen analysis, site context, etc.) can a
truly rounded picture of Australian human pre-
history emerge.

“Major studies now completed or under way
are beginning to explore this past in Western
Australia. In the South-West, Devil’s Lair (9)
provides the longest continuous occupation
sequence as yet studied (Dortch, in press). In the
Western Desert, (Puntutjarpa (23) Gould
1977) examines the special circumstances of

desert adaptation, while in the Kimberley, Dortch
(in Wright 1977) traces human occupation in
another environment. A number of other studies
examine aspects of Western Australian prehis-
tory (e.g. Clarke and Dortch 1977; Hallam 1972,
1973, 1975, 1977).

“While these studies illustrate what might
best be described as a basic unity in economic
adaptation for at least the later stages of the
prehistoric period considered here, they only hint
at those aspects of prehistoric life concerned
with the more human quality of day to day
living. However, traces of this side of prehis-
toric life are to be found in a wide variety of
rock art that survives today (see Ucko 1977 for
a number of Western Australian examples), some
of which may eventually be shown to be of
considerable antiquity. Finally, it is quite likely
that contemporary concepts as to prehistoric
human occupation and adaptation are far too
rigid in ignoring the significant changes in the
prehistoric environment which doubtlessly had
direct bearing on daily living, as well as enhanc-
ing the very human capacity for social and cul-
tural change.”

Discussion and conclusions

If one had to express in just one word the
salient characteristic of the whole Western Aus-
tralian environment at the beginning of the
historic period from a human point of view, an
appropriate word might be “harsh”, “arid” or
“difficult”. Perhaps the Aboriginal people of the
time did not see it this way because they were
inheritors of a very long tradition of coping with
it. The English invaders certainly did see it so.
The exchange in 1838 between the Aboriginal
Imbat and the Englishman Grey, reported by
Hallam (in Wright 1977) is very illuminating on
this subject.

What then would our time traveller, beginning
here and taking this excessively comprehensive
view, notice as he went back through 200 000
years? If the environment was harsh to begin
with, it must have been harsher still from about
10 000 to about 100 000 years ago, forbiddingly so
perhaps about 14 000 years ago. During the
period of generally high sea levels from about
100 000 to about 130 000 years ago, life might
have been a little easier than in the early his-
toric period, but only marginally. Then more
difficult conditions would be the norm again
almost until the arbitrary beginning of our “pre-
historic” period 200 000 years ago. In short, at
this level of generalization, the time traveller
would register changes only in degree, not in
kind, and indeed he might not notice change in
kind until he had travelled backward more than
10 000 000 years. The Western Australian en-
vironmental stage had been set by middle Ter-
tiary time, and changed only in details subse-
quently.

But man, like all living things, makes his
environmental adjustments to the fine detail,
and our time traveller would need to descend
from his grand geological perspective in order
to make sense of the movements he would see.
Thus the Warburton region (23) populated for

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Journal of the Royal Society of Western Australia, Vol. 62, Parts 1-4, 1979.

the last 10 000 years, might have been truly
“desert” previously. The extreme south-west,
on the other hand might have involved its sparse
human population in unremitting battle with
forest, kept at bay only by the skilful use of fire,
for most of the last 10 000 years, whereas it
might have been more open vegetationally and
certainly did support a much greater variety of
game, for more than 30 000 years before that.
At the latitude of Perth, the coastal, especially
riverine, areas might have been relatively densely
populated, the jarrah forest practically empty,
the wandoo woodland populous and the low
woodland occupied, even if only at low density,
well to the east of Kalgoorlie for most of the
last 10 000 years, perhaps with a shift of popula-
tion densities towards a coast which was itself
continually shifting prior to that.

The separate elements of the environment
would be familiar to our hypothetical time
traveller, but he would note differences in their
relationships to one another, particularly in their
spatial relationships, probably most strikingly in
those depending on the evanescent boundary
separating land from sea. In those prehistoric
intervals of particularly rapid rise or fall of sea
level, for example the rise between, say, 15 000
and 7 000 years ago, not only the time traveller,
but also individual people immersed in the en-
vironment, might perceive changes. It may be
surmised, for example, that an old man living in
the extreme south-west (9) might lament the
submergence of rocks or bays from which he used
to catch fish as a boy. It is even possible that a
percipient person living on the mangrove-lined
fringes of the vast savannah that is now the
Joseph Bonaparte Gulf (34) might notice en-
croachment over a few years, since the sea here
advanced over 250 km in its 90 m rise in level
over this period of about 8 000 years.

The biological component of the environment
is an important one, and in fact a case has been
made (Goldring 1969) that “environments are
essentially the products of biological activity”.
Even some apparently geomorphological features
may be biologically determined, at least in part,
as in the case discussed by Rutten (1967) for
extensive modification by beavers of northern
hemisphere valley forms initially of glacial
origin. Man also has his effects, not only on
vegetation through his use of fire (Hallam 1975)
but in many other ways, for example on the age
structure (and hence human food potential) of
marine mollusc populations under intensive
gathering pressure (Swadling 1976).

Thus our time traveller probably would pay
close attention to the biotic environment, and
would certainly perceive changes, familiar plant
species in unfamiliar places or associations fol-
lowing changes in effective rainfall, or cold-water
molluscs in what are now temperate waters, fol-
lowing changes in air and sea surface tempera-
tures, for example. The theme for prehistoric
time would be changes in range and local (or
total) extinction rather than speciation. For
example, Carroll and Galton (1977) suggest that
“an essentially modern lizard fauna has existed
throughout the Tertiary”, and for those two most
ubiquitous prehistoric and historic components

of the Australian mammal fauna, the macropods
and the murids, Archer and Bartholomai (1978)
suggest a middle Tertiary explosive radiation and
a later Tertiary arrival on the continent respec-
tively, so that both groups were well established
and varied before prehistoric time. Their pre-
historic record seems to be one of response by
established species to environmental changes.

The nature and timing of such responses is
best, though still imperfectly, understood for the
extreme south-west (Cape Leeuwin-Cape Nat-
uraliste region, 9). Here monotremes are known
only from deposits in Mammoth Cave and The
Labyrinth, the former known to be and the
latter suspected of being beyond the range of
radiocarbon dating. Eutherians (murids and
bats) seem curiously under-represented in the
Mammoth Cave deposit in view of their promin-
ence in younger deposits, and in any case have
received little study. Marsupials are therefore
the most reliably and comprehensively known
group. With these provisos, present knowledge
of arrivals and extinctions is outlined in Table
2, and tentative reconstructions of some of the
less familiar species given in Figure 2.

Table 2 lists 5 mammal species of about the
same body size as the extant western grey kan-
garoo, or larger, up to Zygomaturus, of about
the size of a Jersey cow, and at least 8 species
of modest size, such as the koala, involved in a
first phase of local extinction. It shows inci-
dentally that the currently fashionable piece of
jargon “megafauna” not only is unnecessary but
misleading, at least in Australia, where few
species ever exceeded man greatly in size, and
many species of much smaller size have been in-
volved in the “megafaunal” phase of extinction.
Yet it does appear to be characteristic of this
phase that larger species suffered more than
smaller; of those of about human size or larger,
only the western grey kangaroo seems to have
survived in the extreme south-west. Among the
smallest species the murids and bats so greatly
outnumber the marsupials and are so poorly
known, that no such generalization seems war-
ranted.

A second phase of local extinction may be two-
fold, with Sarcophilus and possibly Thylacinus
better considered separately from Perameles,
Bettongia lesueur, Lagorchestes, Petrogale, Pseu-
domys albocinereus, P. occidentals, Notomys and
Macroderma. The latter group make up most
of what Baynes et al. (1976) call the “non-forest
mammals”, i.e. species not recorded from the
south-western forests in historic time, but known
elsewhere from more open or lower plant forma-
tions. Sarcophilus and Thylacinus on the other
hand are listed by Baynes et al. as “forest mam-
mals” by analogy with their distribution in Tas-
mania in historic time; not only did they occur
widely in plant formations of lower or more open
character than our south-western forest, but
they also ranged into similar forest. It
is suggested that the south-western extinction of
Thylacinus and perhaps Sarcophilus is related
to the arrival of Canis , whereas the local extinc-
tion of Perameles and the other non-forest taxa
is related to the spread of dense forest at the
expense of open plant formations.

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Table 2

Arrival, local extinction and survival of mammals in the Cape Leeuwin-Cape Naturaliste region in prehistoric

and early historic time.

1. Present in “Mammoth Cave time" (> 37 000 yr B.P.) See Table 1. Man present. (Provisionally, Zygomaturus,
from Skull Cave, Zygomaturus, Protemnodon, Sthenurus occidentalis, S. brownei, Vombatus hacketti and
Phascolarctos from Devil’s Lair, Macropus (titan?) from Strong’s Cave, and Zaglossus from The Labyrinth,
are assigned here).

2. First phase of extinction (time unknown, on present evidence likely to be greater than 37 000 yr B.P.).
Disappearance of: Zygomaturus, Protemnodon brehus, large ?Protemnodon, Sthenurus brownei, S. occiden-
talis, “Mammoth Cave wallaby or wallabies", Thylacoleo, Onychogalea, Vombatus hacketti, Phascolarctos,
Zaglossus, Tachyglossus aculeatus and larger Tachyglossus species.

3. Phase of arrivals, covering several thousand years before and after 30 000 yr B.P. Bettongia penicillata, B.
lesueur, Petrogale and possibly Lagorchestes, Pseudomys occidentalis and Macroderma gigas make their first
appearances.

4. Second phase of extinction. Holocene. Disappearance of: Sarcophilus, Thylacinus, Perameles, Bettongia

lesueur, Lagorchestes (presumably), Petrogale, Pseudomys albocinereus, P. occidentalis, Notomys, Macroderma.

5. Late arrivals: Canis (before the local extinction of Thylacinus; Merrilees 1979), Rattus tunneyi (early Holo-
cene; Porter 1979), Pseudomys praeconis (late Pleistocene; Balme et al. 1978).

6. Survivals from “Mammoth Cave" to historic time: Dasyurus, Phascogale. Antechinus, Sminthopsis, Isoodon,
*Trichosurus. Pseudocheirus, Potorous, Macropus eugenii, M. irma, M. fuliginosus, Setonix, Pseudomys short-
ridgei, Ratty s fuscipes and possibly one or more species of bats. (* Absent from the lower layers in Devil's
Lair, Balme et al. 1978).

7. Survivals from early or middle “Devil’s Lair” to historic time, not represented in Mammoth Cave Pleistocene
deposits: Cercartetus, Tarsipes, Bettongia penicillata, Hydromys and possibly one or more species of bats.

There does not appear to be any other Austra-
lian region for which a prehistoric faunistic
timetable, even one as loosely framed as Table 2,
can be drawn up yet. But it would not be sur-
prising if the Table 2 theme of nett impoverish-
ment were to be established elsewhere, in view
of the richness of some known prehistoric faunas
(e.g. Kangaroo Island, Hope et al. 1977; the
Murray River in South Australia, Wakefield in
Mulvaney et al. 1964; the lower Darling River,
Merrilees 1973a) or some which may or may not
be prehistoric (e.g. Bingara, New South Wales,
Marcus 1976; Gore, Queensland, Bartholomai
1977).

Reasons for the first south-western phase of
impoverishment remain obscure. It is not yet
clear whether it was exactly or approximately
contemporaneous with its counterparts elsewhere
in the southern part of the continent, as a
climatic interpretation might require. Too little
is known of human modifications of vegetation to
judge their impact on mammal species of largely
unknown ecological characteristics. Practically
nothing is known of prehistoric human popula-
tion densities or subsistence patterns, even
whether large mammals were or were not hunted.
Hence it seems premature to postulate major
effects of human hunting on mammal species of
equally unknown population densities.

But it does seem legitimate to infer vegeta-
tional diversity from the known mammal diver-
sity in the extreme south-west, even though this
cannot yet be checked by pollen or other botani-
cal sequences. Our time traveller would pro-
bably notice differences between times of high
and low sea level. Continuous forest would
accompany high sea levels, and a more diverse
vegetational mosaic would accompany low sea

levels, possibly to the extent of restricting karri
or even jarrah and marri to humid refugia such
as dolines or sheltered valleys, while banksia or
other open woodland occupied much of what was
forested in early historic time. Coastal heath
almost certainly would have been much more
extensive during low than high sea levels.

This diversity in times of low sea level was
such that Potorous and Setonix could continue
to flourish, and Hydromys persist for at least
some of the time. It seems unlikely, therefore,
that extreme aridity could account for the first
phase of extinction, and direct or indirect human
effects seem more likely.

Acknowledgements . — I am grateful to G. W. Kendrick
and M. Lofgren for criticism of the text and to the
latter for supplying, at short notice, the review of the
human component of the environment, to M. Wallis
and E. Lawson for typing, and to C. ten Raa for the
preparation of the figures.

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Aspects of Science in Western Australia 1829-1979.

A volume to commemorate the 150th Anniversary of the founding of Western Australia.

Late Quaternary climates of Western Australia:
evidence and mechanisms

by K.-H. Wyrwoll

Geography Department, University of Western Australia, Nedlands, W.A. 6009

Abstract

An attempt is made to provide an outline of the climatic history of Western
Australia over a period of the order of 80 000 years, for which some information
is available even if sketchy and poorly dated. The late Quaternary was characterised
by marked changes in climate, with possibly quite contrasting conditions occurring
simultaneously in different parts of Western Australia; the dominant feature, however
was an intensification of aridity. A massive extension of the arid zone coincided
with the last global glacial maximum. A consideration of the mechanisms responsible
for this event suggests that this was a recurrent theme of the Pleistocene, an inter-
pretation which appears to be supported by the available stratigraphic evidence.

Introduction

In recent years substantial progress has been
made in establishing the Quaternary palaeo-
climatic history of Australia (see Bowler et al.
1976). Unfortunately our knowledge of the
Western Australian palaeoclimatic record is
sparse when compared to its eastern counterpart.
While there exists a large volume of literature
which contains reference to Quaternary climatic
changes that may have taken place in Western
Australia, much of this work suffers from the
absence of a satisfactorily-dated framework
within which the scattered observations can be
placed, and is marred by the tendency of some
authors to invoke climatic change as a ready
explanation of features which in many cases may
be adequately explained by other factors. More
recent work, which has been facilitated by both
the documented climatic changes that are known
to have taken place in other parts of Australia,
and the initial results of the Climap project,
has shown an explicit concern with the nature
and sequence of Quaternary climatic events in
Western Australia. It is largely from this work
that a framework for understanding the Quater-
nary climatic and environmental history of
Western Australia has begun to emerge.

The aim of this paper is (1) to survey the
major evidence that has been advanced by
various authors to reflect the Quaternary
climatic variations that may have taken place in
Western Australia, and from the evidence
attempt to develop an outline of the late Quater-
nary climatic history of Western Australia, (2) to
formulate the mechanism of one palaeoclimatic
event — widespread aridity — which is likely to
have been a recurrent feature of the Quaternary
climatic history of Western Australia. As far as
possible the evidence is reviewed under separate
subject headings, but in places material which is
not encompassed by a particular section heading
is included to avoid clumsy cross-referencing and

repetition between sections. Figure 1 shows the
location of many of the sites referred to in the
text.

Biotic evidence

Evaluating the climatic implications of faunal
and floral remains has been one of the mainstays
of Quaternary palaeoclimatic reconstruction. In
Western Australia few studies of biotic remains
have so far been undertaken with the specific
aim of reconstructing the Quaternary climatic
history of the area. Especially regrettable is the
limited use that has been made of palynology, a
technique which in other parts of the world has
been fundamental in elucidating the Quaternary
environmental history.

Biotic distributions

As probable indicators of Quaternary climatic
events in Western Australia, disjunct distribu-
tions of flora and fauna have attracted a great
deal of attention. Considerable discussion has
surrounded the apparent west-east and north-
south affinities in the distribution patterns of the
Australian flora (Gardner 1942; Specht 1958;
Burbidge 1952, 1959, 1960; Gentilli 1961; Green
1964; Marchant 1973; Nelson 1974). A compari-
son of the vascular floras of south-west and
south-east Australia by Green (1964) revealed
the existence of several hundred species common
in both areas but not in the drier intervening
regions. Green considered that some of these
populations are true vicariads (defined as closely
related allopatric species which have descended
from a common ancestral population and
attained at least spatial separation) which
once occupied continuous areas which became
broken by the onset of drier climatic conditions.
Some examples include Levidosverma angusta-
tum (south-west), L. concavum (south-east);
Casuarina decussata (south-west), C. torulosa

82710 ( 9 )

Figure 1. — Location map showing the major sites mentioned in the text.

130

Journal of the Royal Society of Western Australia, Vol. 62, Parts 1-4, 1979.

(south-east); Acacia leptoneura (south-west), A.
rigens (south-east). Estimates as to when this
separation of the south-west and south-east
flora occurred are widely conflicting, and range
from the late Tertiary to the late Quaternary.

North-south affinities in plants are well docu-
mented in Western Australia. Gardner (1942)
for instance, explained the occurrence in the
Geraldton-Perth area of plants common in the
Kimberley district, (e.g. Cartonema, Dioscorea
and Clematicissus) , as the result of the extension
of tropical conditions further south at some
time in the past. Other examples include the
existence of species of Terminalia, Owenia, Mal-
lotus and Lysiphyllum in the Pilbara district
(Burbidge 1959). The occurrence in tropical
areas of isolated species of genera which are well
developed in the South-West Botanical Province
has been interpreted by several authors as being
indicative of the former extension of southern
climatic conditions further north (Gardner 1942;
Burbidge 1952, 1959, 1960; Gentilli 1961). One
species of Byblis occurs only in the South-West
Province while the only other species in the
genus is widespread in the tropical zone. Verti-
cordia is mostly confined to the South-West
Province and a contiguous semi-arid belt further
inland, while in the tropical zone the genus is
represented by several species. Other examples
include species of Borya, Persoonia, Boronia,
Jacksonia and Calytrix (Burbidge 1960). A relict
plant distribution which has not received much
attention is the Millstream Palm ( Livistona
alfredii ) which is confined to the Fortescue River
area at Millstream and a few nearby localities.
This species represents the only natural occur-
rence of palms in Western Australia outside the
Kimberley district (Environmental Investiga-
tions, Fortescue River 1975).

A number of writers have explained the mod-
ern distribution of some amphibians, reptiles,
marsupials and birds, east and west of the Nul-
larbor Plain as the result of greater aridity in
the past (e.g. Serventy and Whittell 1976; Main
et al. 1958; Keast 1961; Kluge 1967). Main et al.
(1958) explained the existence of frogs with
eastern affinities in south-western Australia as
the result of successive movements from the
primary speciation area in south-eastern Austra-
lia across the southern Nullarbor Plain during
“pluvial” periods of the Pleistocene. Interven-
ing arid periods were seen as the mechanism of
isolation of the eastern and western faunas.
Serventy (1951) reached similar conclusions con-
cerning the speciation of the chestnut-shouldered
wrens (Malurus) which occur throughout the
Australian region. Serventy and Whittell (1976)
noted numerous examples of the discontinuous
occurrence of related avian fauna in Australia.
Again the onset of arid conditions during the
Pleistocene was invoked to explain these features.
Ford (1974), however, has suggested that some
isolates of desert species of birds may have
formed during relatively wet periods in the
Pleistocene.

The only way that the various authors were
able to suggest a possible time at which the dis-
tributions became separated was by placing them

in the context of the palaeoclimatic record
available at their time of writing; as the record
was often incomplete or in error their conclusions
were understandably misleading. This necessity
to resort to an existing palaeoclimatic chron-
ology, in order to date a climatic event, postu-
lated on the basis of a disjunct distribution, is a
major limitation in the use of disjunct distribu-
tions in palaeoclimatic reconstruction. Conse-
quently biotic distributions have been of little
value in building up the chronology of Quater-
nary climatic events in Western Australia,
although they have proven useful in indicating-
likely climatic events that may have taken place.

Nullarbor Plain

Biological evidence for Quaternary climatic
changes in the Nullarbor Plain is sparse and
often poorly dated. Several workers have sug-
gested that the climate of this region has not
changed significantly in the last 20 000 years.
Faunal remains found in sediments in Koonalda
Cave were dated at approximately 20 000 years
BP (Wright 1971, p. 24). Thorne (1971) found no
evidence from these faunal remains suggesting
an environment different from that of today.
Martin (1973) examined pollen from three
excavations — N145 Cave near Eucla, Madura
Cave and Norina rock shelter. She found that
the period 20 000 to about 10 000-8 000 years BP
appeared to be more arid than the climate of
that area today, with annual rainfall averaging
about 180 mm. From about 10 000-8 000 to 5 000-

4 000 years BP rainfall apparently increased, and
has since maintained an annual average of about
250 mm. The climatic changes indicated by the
varying pollen ratios were primarily explained by
Martin as being due to the influence of a rising
sea level and the consequent climatic effects
resulting from a closer proximity to the coast.
Martin also suggested that a change in the
level of the Aboriginal population at about 4 000-
6 000 years BP in the area may have influenced
the vegetation patterns.

There is some additional evidence that seems
to support Martin’s (1973) conclusion that since
the middle Holocene the rainfall received by the
Nullarbor Plain area has been at the same level
as today. The preservation of soft tissue on a
Thylacinus cynocephalus (Tasmanian tiger)
skeleton in Murra-el-elevyn cave (dated at 3 300
years BP, Partridge 1967) and in Thylacine Hole
(dated at 4 600 years BP, Lowry and Merrilees
1969) suggested to Lowry and Jennings (1974)
that conditions in the caves could not have been
much wetter in the intervening time than at
present. Ingram (1969) examined the pollen
grains found in the desiccated gut of mammalian
carcases — Megaleia rufa (red kangaroo), Tricho-
surus vulpecula (possum), Onychogalea lunata
(crescent nail-tail wallaby) and Canis familiaris
dingo (dingo) — found in Thylacine Hole. These
carcases are considered to be between 2 000 and

5 000 years old because they were associated with
a Thylacinus cynocephalus carcase which has
been radiocarbon dated at 4 650 ± 153 years BP,
and a Canis familiaris dingo carcase which has
been dated at 2 200 ± 96 years BP (Lowry and

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Journal of the Royal Society of Western Australia, Vol. 62, Parts 1-4, 1979.

Merrilees 1969). The fact that the fossil pollen
belong to genera common in the present day
vegetation, led Ingram (1969) to conclude that
the climatic conditions in the area 2 000 to 5 000
years ago, were similar to those of today.

The conclusions of Ingram (1969), Thorne
(1971) and Martin (1973) support the view of
Jennings (1967), based on the retarded karst
development of the Nullarbor Limestone, that
there is little evidence for significant climatic
changes in the Nullarbor Plain during the Quat-
ernary. In a more recent publication Lowry and
Jennings (1974) suggested that there have been
changes in the water availability and effective
precipitation in the Nullarbor karst during the
Quaternary, but that they have never departed
substantially in either direction from the present
state. They concluded that the karst develop-
ment on the Nullarbor “may owe much to the
cold phases of the Pleistocene when effective
precipitation, if not absolute precipitation, was
in all probability greater than at present though
still low” (Lowry and Jennings 1974, p. 79).

There is some evidence which may support the
view of Lowry and Jennings that there was an
increase in effective precipitation on the Nullar-
bor Plain during the Pleistocene glacial periods.
Glauert (1912) found the remains of relatively
large extinct marsupial species in the Balladonia
area, and interpreted them as evidence of a more
moist climate at some time during the Pleisto-
cene. Similarly the finding of Sthenurus (an
extinct large kangaroo) in Madura Cave led
Lundelius (1963) to infer more humid conditions
in the Pleistocene. These arguments for wetter
conditions were based on the assumption that
the “giant” marsupials could not have survived
in relatively dry habitats. Such an interpreta-
tion can be questioned because existing large
marsupials in Australia appear to thrive under
conditions more harsh than those now prevailing
at the fossil sites of the Nullarbor Plain (Lowry
and Jennings 1974). However, other evidence
which may support Glauert’s conclusion, is
offered by Richards (1971) who considered that
the ancestors of various arthropods found in
Nullarbor caves must have colonised that area
when conditions were wetter. Furthermore,
fossil land snails found in kankar from the
Nyanga Plain (the area to the north and west
of the Nullarbor Plain) were tentatively com-
pared with the living species Bothriembryon
barretti by G. Kendrick (in Lowry 1970). While
the fossil snails were found within the 150 mm
isohyet, Lowry (1970) has found living snails
only in areas with more than 250 mm of rain-
fall. With the absence of absolute dates and
reliable stratigraphic information it is not pos-
sible to use this evidence to evaluate the sug-
gestion that the climate of the Nullarbor may
have been wetter during the glacial periods of
the Pleistocene. Nevertheless, the evidence does
at least suggest that wetter conditions have
occurred.

South-west

Early work on cave deposits in the Cape
Leeuwin-Cape Naturaliste region by Glauert
(1910, 1914), Lundelius (1960) and others has

been revised in a series of papers by Merrilees
(1968a, b), Dortch and Merrilees (1972), Baynes
et al. (1975), Balme et al. (1978), Porter (1979)
and Merrilees (1979). They report a well-dated
continuous sequence beginning before 37 000
years BP (Mammoth Cave), from about 35 000
to 5 000 years BP (Devil’s Lair) and from before
8 000 years BP to the present day (Skull Cave),
with notes on other less well-dated sites.

It is clear that the region supported a very
diverse mammal fauna in the Pleistocene. Some
of the species represented in the cave sequences
have a wide climatic tolerance (e.g. Trichosurus
vulpecula — brush-tailed possum) ; species indica-
tive of drier conditions are also found (e.g. Bct-
tongia lesueur — burrowing rat-kangaroo) ; and
remains of species which required rather wet,
well-vegetated surroundings are also present in
the cave deposits (e.g. Potorous tridactylus —
long-nosed rat-kangaroo). At some time not yet
precisely dated (or perhaps over some consider-
able period) many large species including Zygo-
maturus trilobus (a diprotodontid) disappeared
from the record, and at some time later (late
Pleistocene to early Holocene) several of the
species most characteristic of drier climates, in-
cluding Macroderma gigas (ghost bat), retracted
their ranges out of the region.

The reasons for the disappearance of the large
extinct species are still obscure, but Merrilees
(1979) suggests that the “dry climate” species
were displaced in early Holocene time by the
development or extension of dense karri forest
like that of early historic time. The implication
is that higher effective rainfall, perhaps with less
extreme difference between wet and dry seasons,
marked the beginning of the Holocene.

An indication of possible Holocene climatic
variations in the extreme south-west is given by
Churchill (1968), who suggested that rainfall
is the major determinant of the distribution of
Eucalyptus marginata, E. diversicolor and E.
calophylla. Changes in the E. diversicolor /E.
calophylla ratio were thought to provide an index
of precipitation changes; a relative increase of
the E. diversicolor / E . calophylla ratio being
indicative of wetter conditions. It appears from
pollen analyses at Flinders Bay Swamp, Weld
Swamp and Boggy Lake that conditions were
relatively wetter between 6 000 and 5 000 years
BP, followed by a trend towards drier conditions
until about 2 500 years BP; after which wetter
conditions apparently prevailed until about 1 300
years BP, subsequently it became drier until
about 500 years BP. From then to the present
there was a trend towards higher rainfall. The
similarity of Churchill’s findings, that in the
south-west conditions were wetter between 6 000
and 5 000 years BP, with the changes recorded
in lakes of western Victoria (Dodson 1974a, b)
and on Wilson’s Promontory, eastern Victoria
(Hope 1974) led Bowler et al. (1976) to suggest
that this constituted a climatic event which
affected much of southern Australia. Kendrick
(1978) described an assemblage of land snails
from palaeosols occurring on and beneath the
surface of an elevated ridge of aeolian calcar-
enite from near Point d’Entrecasteaux. He con-
concluded that the assemblage — Bothriembryon

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Journal of the Royal Society of Western Australia, Vol. 62, Parts 1-4, 1979.

gardneri, B. consors, species of Paralaoma, Magi-
laoma, Pernagera, one other charopid and an
undescribed assimineid — suggests a humid, well
vegetated, probably forested environment in the
Point d’Entrecasteaux area at the time of soil
formation, in contrast to the exposed coastal
heath that presently characterises the area.
Unfortunately the actual age of this assemblage
is as yet unknown.

Kendrick (1977) found that fossil molluscs
from a mid-Holocene deposit near Guildford
have a greater marine affinity than the present
assemblage in that area. The fossil deposits
have a radiocarbon age of 6 600 ± 120 years BP,
coinciding approximately with the Holocene mar-
ine transgression. It follows that a greater mar-
ine influence would be expected in this part of
the Swan River at that time. But despite this,
given the present river discharge conditions near
Guildford, Kendrick argued that river discharge
and seasonality were much lower than at present,
which implies that drier climatic conditions pre-
vailed in the Guildford region from at least 6 700
years BP until some time after 4 500 years BP.

Northern areas

Kendrick (1978) has recently described two
new species of Bothriembryon snails from pal-
aeosols in the Shark Bay district. The two
species, B. douglasi and B. ridei, have no known
living descendant in that area. Their extinction
at about 80 000 years BP, and the subsea.uent
appearance of camaenid snails ( Rhagada , Plec-
torhagada and Angasella species), as well as the
size reduction of B. costulatus, are thought to
have resulted from the onset of more arid con-
ditions in that area at that time. More humid
conditions are thought to have returned to the
area since about 10 000 years BP.

Other evidence of Pleistocene climatic events
in northern Western Australia comes from Mayr
(1944) who discussed the avian faunal exchanges
which evidently occurred between Timor and
Australia during the period of each Pleistocene
glaciation. The fact that most of the recent
arrivals from Timor were grassland birds led
Mayr to conclude that the climatic conditions
prevailing during each glacial were at least as
dry or drier than at present. A vertebra from
the Ederga River and a tooth from the Gascoyne
River district mentioned by Merrilees (1968b, p.
15, the latter tentatively classed as “varanid”)
may represent crocodilians (D. Merrilees, 1978,
pers. comm.), and if so, would imply a possible
southward extension of tropical conditions at
some time in the Pleistocene.

Gorter and Nicoll (1978) have recently dis-
cussed the occurrence, at Windjana Gorge, of
indeterminate crocodilian and turtle remains in
river gravels of possible Pleistocene age, which
may however be as old as early Miocene. The
authors claim that the crocodilian remains do
not have any important palaeoenvironmental
significance. But the presence of remains of the
river turtle Carettochelys implies a warmer
climate than the present one in north-western

Australia. However, Gorter and Nicoll stress
that without more evidence this conclusion is
speculative.

An indication of part of the Holocene climatic
history of the northern areas is given by Jen-
nings’ (1975) work in the Fitzroy estuary. At
present this area experiences a long dry season
which in part results in low mangrove forest and
mangrove scrub. The stratigraphy of two
embayments in the Fitzroy estuary indicates a
much wider extent of mangrove swamp at about
7 400-6 000 years BP. In addition the occurrence
of fossil stumps of mangrove larger than those
of today suggest that tall mangrove forest pre-
vailed at that time. From this, Jennings con-
cluded that at the time a longer and heavier wet
season characterised the Fitzroy region. In sup-
port of his conclusions Jennings cited Stocker
(1971) who suggested that a decline in annual
rainfall or a lengthening of the dry season had
taken place on Melville Island between 8 000 and
2 000 years BP. Present day jungle fowl on
Melville Island inhabit closed communities, par-
ticularly monsoon forest. The discovery of
ancient mound nests in eucalypt forest led
Stocker (1971) to suggest that there has been a
gradual decrease in monsoon forest on the island
resulting from either a reduction in rainfall or
the effects of the Aborigines through their use
of fire.

Geomorphological-geological evidence

Longitudinal sand dunes occupy a large area
of the Australian continent, but are presently
active only in the arid centre. The occurrence
of vegetated dunes on the present desert margins,
similar in form to their more arid counterparts,
provide the most striking evidence of the exten-
sion of more arid conditions, at some time in the
past. An understanding of the timing and dev-
elopment of these sand dunes, taken in conjunc-
tion with the evidence from lake lunettes, has
added much to our understanding of Australian
Quaternary palaeoclimates (see Bowler (1976)
for a review of much of this evidence).

In Western Australia aeolian evidence for a
former extension of arid conditions is wide-
spread. Veevers and Wells (1961) described the
now stabilised sand dunes and sheets of the
Canning Basin, and attributed their development
to an arid phase some time in the Pleistocene.
Earlier work by Fairbridge (1953) and Brunn-
schweiler (1957) had already led to the recogni-
tion of now stable large aeolian bedforms in this
area. Both authors attributed them to the oc-
currence of extended arid conditions during the
Pleistocene. Brunnschweiler (1957) suggested
that the inland dunes of the Dampier Peninsula
(a term he applied to the area west of King
Sound) are continuous with dunes further in-
land, and proposed that the dunes formed in the
Pleistocene during a period corresponding with
the Riss Glacial of Europe; however there is no
evidence put forward by the author which sup-
ports this age estimate. Fairbridge (1953) fol-
lowed Mayr’s (1944) discussion, referred to in an
earlier section, and concluded that these dunes
stem from a glacial period. In a later work

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Journal of the Royal Society of Western Australia, Vol. 62, Parts 1-4, 1979.

Fairbridge (1964) compared the way the fixed
longitudinal dunes in the Fitzroy area appear to
descend below present sea level, as also described
by Wright (1964), with an analogous situation
in the lower Senegal Valley, and postulated arid
conditions and associated dune formation in the
Fitzroy area during the last glacial.

The stratigraphic evidence provided by Jen-
nings (1975) confirmed the superficial appearance
of the burial of the desert dunes by estuarine
fill. A minimum age of 8 000 years BP was in-
ferred from the depth of buried dune sand, while
a maximum age of about 140 000 years BP was
suggested on the basis of well-presented dune
shape. Jennings (1975) related his findings to
those of van Andel et al. (1967) who interpreted
the occurrence of brown calcareous nodules
(kankar) in the sediments dated at 17 000 years
BP on the Sahul Shelf, as evidence of drier
climatic conditions at the last glacial maximum.
From this evidence Jennings (1975) reached the
conclusion that the most probable time for dune
formation in the Fitzroy area was at approxi-
mately 20 000 years BP.

Evidence in support of this age estimate for
the formation of these dunes comes from the
abundance of quartz in Indian Ocean sediments
dated at about 18 000 years BP (Kolia and Bis-
caye 1977). The quartz particles found in these
sediments are derived from the adjacent contin-
ental area, and the higher amounts of quartz in
the sediments suggest increased continental
aridity and/or increased intensity of wind at the
time of deposition.

Further south in the Exmouth Gulf area at
least two inland dune units can be distinguished
(Wyrwoll, unpublished). Both appear to predate
the Holocene transgression. The older dune
unit is capped by a thick indurated calcrete
horizon, while on the younger dunes more weakly
developed calcareous soils are found. No abso-
lute ages for these dune units are yet available.
The presently vegetated dunes of the Gascoyne-
Minilya-Lyndon Plain appear to correspond to
the younger dunes of the Exmouth Gulf. This
dune unit seems to predate the evaporite
sequence of Lake Macleod. So far it has not
been possible to develop a convincing strati-
graphy of the Quaternary sediments of the Ex-
mouth Gulf -Gascoyne area which would time
bracket the two dune units with sufficient resolu-
tion.

Logan et al. (1970) describe two red sand dune
units in the Shark Bay region. In the upper
part, the older unit (the Peron Sandstone) has
a strongly developed nodular calcrete horizon.
The Peron Sandstone may correspond to the
older dune unit recognised in the Exmouth Gulf
area. Although Logan et al. provide a compre-
hensive account of the stratigraphy of the Quat-
ernary sequence of the Shark Bay area, the abso-
lute age of the Peron Sandstone remains uncer-
tain. However, the Peron Sandstone is overlain
by the Dampier Transgression, and 2;!0 Th/ 234 U
dating on corals from this transgression is now
being undertaken (W. J. E. van de Graaff, 1978,
pers. comm.), which will provide at least a
minimum age for the Peron Sandstone. The

younger dune unit, the Nilemah Sands of Logan
et al., consists of unconsolidated red sands which
overlie the other Pleistocene formations of this
area. This unit corresponds to the unconsoli-
dated dunes of the Gascoyne Plain-Exmouth
Gulf region. Degraded inland dune forms can
be identified as far south as the Geraldton-
Mullewa area (Johnson et al. 1954; Wyrwoll
unpublished). Further inland Mabbutt (1963a,
b) found that an arid event affected the Murchi-
sonia and Salinaland divisions (Jutson 1950),
but was unable to determine an age for this
event, except to conclude that it took place some
time during the Pleistocene. The yellow sands
which occur throughout the Perth Basin have
been interpreted by Glassford and Killigrew
(1976) as a relict aeolian desert sediment derived
from sources on the Yilgarn Block and in the
Perth Basin. Their conclusion is based on the
textural characteristics and mineralogy of the
sands, and the occurrence in the sands of kaolin
spherites (see Killigrew and Glassford 1976)
which they claim have not been observed in
marine derived coastal dune or aeolian calcar-
enite sediments.

In the south Bowler (1976) has dated lunette
deposits which fringe Lake Kurrenkutten,
Storey’s Lake, Lake Grace and Lake King. The
results indicate that a short-lived lunette-
building event took place between 20 000 and
15 000 years BP, during a period of hydrologic
stress; this is an event which has been widely
recognised throughout much of southern Aus-
tralia.

Other geomorphological evidence of Quater-
nary climatic change in Western Australia is
not well documented. The conclusions that Jen-
nings (1967) and Lowry and Jennings (1974)
arrived at with respect to the degree of karst
development on the Nullarbor Plain have already
been noted. Wright (1964) envisaged that the
onset of drier climatic conditions in the Pleisto-
cene led to the formation of floodplains along
the Fitzroy and Lennard Rivers and their tribu-
taries; and to the extensive alluviation mar-
ginal to upland areas in the Fitzroy Basin.
Further south in the Geraldton area a “Red
Alluvium” unit has been described, and dated
at earlier than approximately 40 000 years BP
(Wyrwoll 1977). The “Red Alluvium” overlies
marine deposits, which on the basis of the mol-
luscan fauna and coastal stratigraphy, were
attributed to a last interglacial high sea level,
thought to have occurred at about 120 000 years
BP (Veeh 1966). At the time of deposition of
the “Red Alluvium” streams carried a much
coarser bed load than at present, and appear to
have had a greater competence and capacity
than the present streams. Sediment yields at
the time of deposition of the “Red Alluvium”
were almost certainly considerably higher than
at present. It was tentatively concluded that
semi-arid conditions with intense rainfall events
occurring more frequently than today, may have
prevailed at the time of deposition of the “Red
Alluvium”.

A fine textured, inset alluvial fill from the
Geraldton area, described informally as the
“Grey Alluvium” and originally dated at 3 185

134

Journal of the Royal Society of Western Australia, Vol. 62, Parts 1-4, 1979.

± 145 years BP (Wyrwoll 1977) has recently
been used by Williams (1978) to support a postu-
lated continent-wide climatic change between
approximately 4 000 and 1 500 years BP. In the
Southern Tablelands of New South Wales this
event may have led to a change in rainfall
seasonality, to lower temperatures, and to drier,
windier conditions. Since the original date was
published an additional 14 C date with an age of
1 375 ± 120 years BP (GX-5133), has been
obtained for the “Grey Alluvium”. The two
dates demonstrate the diachronous nature of this
alluvial deposit, and question the validity of
using the older date as a reliable upper limit for
the age of the “Grey Alluvium”.

A discussion of the palaeoclimatic significance
of palaeosols found in the Tamala Limestone
was undertaken by Fairbridge and Teichert
(1952). These authors believed that the soil
horizons which separate a number of dune
generations in the Tamala Limestone indicate
minor and short-lived climatic cycles. They
claim a change from rendzinas near Hamelin
Bay, to terra rossas at the latitude of Fremantle,
and travertine crusts along the coast, further
north of Houtman Abrolhos. The authors sug-
gest that this gradation of fossil soils confirms
their belief that at times during the Pleistocene
most of the south-west was subject to more arid
conditions.

Limitations of the evidence

Several authors have pointed out the limita-
tions of the various lines of evidence that have
been used for Quaternary palaeoclimatic recon-
struction in Australia (Jones 1968; Merrilees
1968b; Ride 1968; Calaby 1971; Galloway 1971;
Walker 1976, 1978). It has become apparent
that palaeoclimatic inferences drawn from plant
and animal remains should be made with cau-
tion. Walker (1976) has painted a rather grim
picture of the present reliability of pollen data
for postulating past climatic conditions. In the
context of faunal remains and distributions
Calaby (1971) has pointed out that homoiother-
mic and at least the larger poikilothermic
animals, have a wide climatic tolerance. The
possible role that the Aborigines played in modi-
fying the ecology of the Australian biota (Tin-
dale 1957; Merrilees 1968b; Jones 1968, 1975;
Hallam 1975) must also be considered when
interpreting biotic evidence in terms of palaeo-
climatic events.

Similarly the geomorphological-geological evi-
dence poses considerable problems. The use of
alluvial deposits as palaeoclimatic indicators has
led to imprecise and often widely conflicting
results (see the summary in Flint 1971, p. 306).
The reasons why such discouraging results have
been obtained can be readily understood when
the factors controlling the likely alluvial response
to climatic changes are considered, and the
equifinality of the problem appreciated (e.g.
Schumm 1965; Gessler 1971). Approaching the
study of alluvial deposits from the point of view
of the palaeohydraulics of the sediments may
overcome at least some of the difficulties.

The absence of reliable dates for dune-building
phases is also a major stumbling block, although
it is hoped that dates from pedogenic carbonate
may at least give some indication of the timing
of these events. The interpretation of the Perth
Basin yellow sand as a relict aeolian desert sedi-
ment has been criticised by Lowry (1977), and
although some of the criticism has been answered
by Glassford and Killigrew (1977), the need for
a more comprehensive presentation and discus-
sion of the evidence remains (cf. Clark and
Dortch 1977, who also consider the age and
origin of the yellow sand).

Chronology of events

The weakness of the available evidence is high-
lighted when any attempt is made to outline the
chronology of the late Quaternary climatic his-
tory of Western Australia. The absence of a
continuous record, the few absolute dates, and
the problems associated with the interpretation
of the available evidence, make it impossible to
give anything other than a crude and incomplete
outline of the late Quaternary climatic history
of this area.

80 000-20 000 years BP. Evidence from the
central coastal area suggests that in this area the
beginning of this period may have seen the onset
of arid conditions, which appear to have pre-
vailed until the beginning of the Holocene. The
early part of this period saw the frequent occur-
rence of high precipitation events leading to high
sediment yields, and resulting in extensive
aggradation in the Geraldton area. The now
drier parts of the south may have experienced an
increase in effective precipitation during part of
this period.

20 000-10 000 years BP. Until about 14 000
years BP intense widespread aridity affected
much of Western Australia. The now stabilised
and partly degraded dune fields, which in the
coastal region extend from the Fitzroy to the
Geraldton-Mullewa areas, were formed during
this period. The Devil’s Lair sequence suggests
a significant reduction in the effective precipita-
tion of the lower south-west at the same time.
In the south-west and Nullarbor Plain relatively
dry conditions seem to have prevailed until the
end of the Pleistocene.

10 000 years BP to present. The climatic
record for this period as for the earliest period
is very incomplete. The onset of the Holocene,
with a rising sea level and higher sea surface
temperature is likely to have seen a significant
increase in precipitation over much of Western
Australia. During the period 7 400-6 000 years
BP, the north-west appears to have received a
higher rainfall which extended further south
than at present. More arid conditions may have
prevailed in the south-west from 6 700 years BP
to about 4 500 years BP. The pollen evidence
from the extreme south-west suggests increased
precipitation over the period 6 000-5 000 years
BP, followed by drier conditions which prevailed
until 2 500 years BP.

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Journal of the Royal Society of Western Australia, Vol. 62, Parts 1-4, 1979.

Mechanism of arid-zone extension

A dominant theme which emerges from the
available literature is the apparent expansion of
the arid zone, probably at various times during
the Quaternary. The most recent widespread
expansion appears to be well documented and
seems to have coincided with the last glacial
maximum (Jennings 1975; Bowler 1976; Wyrwoll
and Milton 1976; Kolia and Biscaye 1977). This
was an event which in Australia led to an exten-
sion of the arid zone along the whole of its
present perimeter (Bowler et al. 1976) ; and which
can be related to the expansion, at that time,
of low latitude arid areas throughout the world
(Sarnthein 1978).

Few attempts have been made to interpret
this generally accepted expansion of the arid zone
in terms of the climatic controls that may have
prevailed at that time. One approach that can
be taken to obtain an indication of the dynamic
palaeoclimatology of past events, is to use the
extremes of the present circulation as an anal-
ogue for past conditions (see Barry 1975). The
assumption inherent in any analogue approach
is that the present-day climatic variability in-
cludes conditions which were sufficiently frequent
in the past to give rise to a substantially different
climatic regime (Dzerdzeevski 1963). This
amounts to assuming a form of climatic uni-
formitarianism which, in the light of changing
boundary conditions and a possibly almost
“intransitive atmosphere”— where the set of
equations which represent the behaviour of the
atmosphere has a number of solutions, each
representing a possible condition of the atmo-
sphere (Lorenz 1976) — is unrealistic. Even over
decades, changes of climatic characteristics have
been recognised that cannot be attributed to
changes in the frequency of a particular circula-
tion pattern, but which arise through changes
in the characteristics associated with the circu-
lation (e.g. Perry and Barry 1973). Despite
these serious limitations an analogue approach
may serve as a useful first approximation to a
partial explanation of past climatic events.

The present climate of Western Australia is
strongly influenced by the subsiding air of the
sub-tropical high pressure belt (STH) (Gentilli
1971, p. 53) which is linked with the descending
branch of the low-latitude Hadley cell— the
direct cause of much of the widespread aridity
of sub-tropical latitudes (Hare 1961). The sea-
sonal displacement of this belt is an important
control on both the summer precipitation of the
north and the winter rainfall of the southern
areas of Western Australia.

The seasonal shifts of the Intertropical Con-
vergence Zone (ITCZ) dominate the dynamic
climatology of the northern areas. It is in this
zone that the tropical rain-producing disturb-
ances originate (Charney 1968; Bates 1970), so
that the location of this feature is fundamental
m determining the rainfall of adjacent areas.
The location of the ITCZ is linked to the meri-
dional temperature gradient of both hemispheres
(Newell 1973), and seems to be determined by
low-level convergence in the tropical boundary
layer (Charney 1968). Prevailing sea-surface

temperatures also seem to play an important role
in determining the location of the ITCZ (Char-
ney 1968; Pike 1971). The ITCZ only reaches
northern Australia in summer when the STH has
been displaced to the south. Associated with the
ITCZ is an inflow of moist, unstable air which
gives rise to the “Australian summer monsoon”.
This monsoon circulation however, is shallow and
weak, and only a pale imitation of its Asian
counterpart.

Despite the fact that during the summer
months the STH cells are located far to the
south, they can still exert a marked influence
on the precipitation of the northern areas. In-
tense anticyclones situated over the Australian
Bight can give rise to a quite stable atmosphere
above the Pilbara heat low, and through this
prevent the incursion of monsoonal depressions
and disturbances into the northern areas
(Rutherford and Hannan 1956; Falls 1970).

A comparison between the rainfall received by
northern Australia in February 1952, and Febru-
ary 1953, highlights the control that the STH
exerts on the rainfall of the northern areas. In
February 1952, when northern Australia was in
the middle of its worst drought in 60 years, the
highs migrated 5° closer to the equator than
normal. In February 1953, however, when rain-
fall was high the track of the migrating highs
lay up to 10° poleward of normal (Ramage 1971.
p. 273).

The winter rainfall of the southern area of
Western Australia is largely associated with
moist unstable westerly winds and with the
associated troughs in this westerly airstream.
The mean depression track and the associated
zone of maximum rainfall are situated off the
south-west coast, and this accounts for the
decreasing precipitation gradient northward
(Wright 1974). Pittock (1973, 1975) has demon-
strated a correlation between the latitude of the
STH and the amount of precipitation received by
the south-west. This correlation has been sub-
stantiated by Wright (1974) whose work focuses
on the south-west of Western Australia. These
authors have shown that a more southerly loca-
tion of the STH can significantly reduce the
winter rainfall received by the south-west.

Using the present climatic regime as an ana-
logue for past conditions it is evident that an
increase in anticyclonicity would directly result
in widespread aridity over much of Western
Australia, Wyrwoll and Milton (1976) have
argued that the lowering of the mean July sea
and land surface temperatures (Climap 1976;
Gates 1976a, b) would have thermally intensified
the anticyclone. The resultant anticyclone would
have combined the high pressure qualities of
both anticyclonic types — warm and cold (Wexler
1951) — and therefore would have been extremely
intense (also see Derbyshire 1971). Under these
conditions the frequency of breakdown of anti-
cyclonic cells would have been considerably
lower, and with the consequent development of
blocking conditions, winter rainfall would have
been confined to the extreme south-west of West-
ern Australia.

136

Journal of the Royal Society of Western Australia, Vol. 62, Parts 1-4, 1979.

To describe the synoptic conditions governing
the summer months during full glacial periods
it was envisaged (Wyrwoll and Milton 1976) that
anticyclonic conditions would have weakened,
but possibly only to the extent that conditions
were more similar to those of the present day
late spring or early autumn. This would imply
that during the full glacial summer the STH
was located significantly further north than
during the present summer months. From the
present day situation it is clear that this would
have been a very effective mechanism in reduc-
ing the rainfall received by the northern areas.

A northward displacement of the STH during
glacial summers is supported by the so called “Z”
criterion developed by Smagorinsky (1963), and
subsequently used by Flohn (1964, 1965) for
locating the position of the STH. The “Z” cri-
terion takes the form

a 91n<9 PE /dy

cot <p = (1)

9 In 0 PE /9z

where 0 is the latitude of the STH, h a scale
height of the atmosphere of about 9 km, which
approximates the height of the 500 mb surface,
and a is the radius of the earth, 0 PE is the partial
potential equivalent temperature (see Appendix
1), and z and y are the vertical and horizontal
axes respectively. Instead of using the potential
equivalent temperature Flohn (1965) has
approximated equation (1) by using the poten-
tial temperature (e) and by relating this to the
actual temperature, Flohn has prepared a nomo-
gram from which the effect of changes in the
meridional and vertical temperature gradients
on the position of the STH can be read (Fig. 2).

0° 2° 4° 6° 8° 10° 12° 14°/ IOOO km

9T/ 9y r-

Figure 2 .— Location of the subtropical high pressure
cell ( 0 ) as a function of the meridional and verti-
cal temperature gradients (after Flohn 1965).

The validity of the “Z” criterion appears to
have been established by Korff and Flohn (1969)
who demonstrated a close relationship between
the monthly mean equator-pole potential tem-
perature gradient — which they took as a crude
approximation for dO PE /9 y — and the position of
the STH. Further support was offered by Pit-
tock (1973), who found that equation (1) fairly
accurately predicts the monthly mean latitude of
the STH over Australia. However, Greenhut
(1977) has recently developed a new criterion
for locating the STH which predicts an
opposite response to changes in 9e PE /dy and
de PE /dz to those predicted by the “Z” criterion.

It is difficult to reconcile the prediction of this
new criterion with the success met by Korff and
Flohn and Pittock when using the “Z” criterion.
These two studies would appear to suggest that
the adoption of the “Z” criterion is a valid first
approximation despite Greenhut’s conclusions.

During glacial periods the overall meridional
temperature gradient of the upper troposphere
may have been similar or even less than that of
today (Kraus 1973; Williams 1978). However,
it is likely that a strengthened meridional tem-
perature gradient occurred in the middle tropo-
sphere over the mid-latitudes. According to
Korff and Flohn these are the latitudes which
determine the location of the STH. From Figure
2 it is evident that an increase in the mid-lati-
tude meridional temperature gradient would
be further enhanced if an increase in the vertical
temperature gradient had taken place. For
equatorial areas Kraus (1973) suggested that
during glacial conditions a three-fold amplifica-
tion of the changes in sea surface temperature
occurred by the time the ascending air reached
the upper troposphere. This occurred as the
result of a decrease in latent heat release, and
would only be significant over areas with a sea
surface temperature greater than approximately
20°C. In addition there is some evidence from
South-East Asia (Verstappen 1975) and New
Mexico (Wright et al. 1973) that during full
glacial conditions a lapse rate of temperature
with height greater than present, may have pre-
vailed at least in the lower latitudes. The com-
bined evidence suggests that an equatorward dis-
placement of the STH during full glacial condi-
tions would have been highly likely.

A northward displacement of the STH should
result in a concomitant displacement of the
ITCZ and the depressions associated with it.
Available evidence (Bryson 1974; Beer et al.
1977) suggests that the amount of northward
displacement of the STH would be amplified,
possibly up to three times, with respect to the
ITCZ. Evidently an equatorward displacement of
the STH would be very effective in causing the
extension of arid conditions in the northern
areas.

In addition, because of the high contribution
that tropical cyclones make to the annual pre-
cipitation of the north-west (Milton 1978), wide-
spread aridity in that area during the last glacial
maximum implies an absence of tropical cyclones
at that time. Whether this was due to an ab-
sence of cyclogenesis as a result of reduced sea
surface temperatures (Webster and Streten 1972)
or simply a more westerly seaward path is not
possible to establish. As tropical cyclones are
very effective converters of potential to kinetic
energy, and play an important role in the general
circulation of the atmosphere (Landsberg 1960),
a possible absence of tropical cyclones during full
glacial periods would have important repercus-
sions on the general circulation.

An equatorward displacement of the STH dur-
ing full glacial stages has also recently been
suggested by Rognon and Williams (1977) and
appears to be supported by the available geo-
morphological evidence. The orientation of

Journal of the Royal Society of Western Australia, Vol. 62, Parts 1-4, 1979.

clay lunettes in southern Australia suggests that
the summer anticyclone over Australia in full
glacial times was positioned some 5° further
north than today (Bowler 1976). In addition
Bowler envisaged a strengthening of the west-
erly air flow over southern Australia at that time.
In Western Australia the geomorphological evi-
dence of dune and lunette orientation appears to
support this reconstruction. But this circulation
pattern is fundamentally different from that
required by Glassford and Killigrew (1976) to
account for the derivation of the Perth Basin
yellow sand. Their interpretation implies that
surface easterlies prevailing as far south as the
Leeuwin Block, occurred with sufficient fre-
quency and intensity to transport large volumes
of sand westward. This, taken in conjunction
with the proposed extension of desert conditions
to the coast, could only occur under a circulation
pattern in which the STH during a large part of
the winter, is located further south than its pre-
sent summer location. From equation (1) it
follows that this could only occur during periods
in which a substantial increase in dd PE /dz and/or
a decrease in dd PE /dy had taken place. While it
would seem that such an extreme southward dis-
placement of the winter STH is unlikely, if it did
occur it would almost certainly have occurred
during interglacial stages.

During the present interglacial the maximum
southward displacement of the STH should have
occurred at the “climatic optimum” (approxi-
mately 6 000 years BP) when increased global
temperatures and higher atmospheric moisture
content may have significantly reduced dd PE /d y
and/or increased d 0 PE /dz. From the previous
discussion of the present precipitation controls,
it follows that if the STH was displaced south-
ward at approximately 6 000 years BP, climatic
conditions more arid than at present should haye
occurred in the south-west of Western Australia
at that time. Kendrick’s (1977) conclusions
seem to support this idea, but this is rather scant
evidence in view of the earlier work by Churchill
(1968), and the discussion of the Australian
mid-Holocene climatic evidence by Bowler et al.
(1976). However, if it is valid that the location
of the STH is as important a control of rainfall
in Western Australia as has been suggested, then
there is an additional way in which the idea that
there was a significant southward displacement
of the STH at about 6 000 years BP can be tested,
and Kendrick’s conclusions at least partially
checked. It follows that if a significant south-
ward displacement of the STH took place at that
time, and that this displacement also character-
ised the circulation of the summer months,
higher summer rainfall extending further south
can be expected in the northern areas at that
time. Jennings’ (1975) conclusion, that during
the period 7 400-6 000 years BP a heavier wet
season characterised the Fitzroy region, may lend
some support for the idea outlined above.

If it is tentatively concluded that during inter-
glacial stages, a significant southward displace-
ment of the STH can occur, which may lead to
more arid conditions in some parts of the south-
west, it remains to be shown whether during
previous interglacial stages, this southward dis-

placement could have been extreme enough to
lead to an extension of arid conditions as far
south as the Leeuwin Block. The suggestion that
such a massive southward extension of arid con-
ditions may have occurred during a past inter-
glacial stage is certainly difficult to reconcile
with what is thought to happen in low-latitude
arid areas during interglacial stages (see
Sarnthein 1978). In addition aridity extending
as far as the coast in a western littoral during
a period in which sea surface temperatures were
higher than at present is difficult to visualise.

Doubts remain whether it is really necessary
to invoke a significant displacement of the STH
to account for the aridity of the northern areas
during full glacial conditions. Manabe and
Hahn (1977) showed that tropical continental
aridity at glacial maximum is closely related to
stronger surface outflow (or weaker inflow) over
tropical areas. This resulted from the hydro-
static response to the greater reduction of sur-
face temperatures over continents than oceans
and the increased albedo value that character-
ised tropical areas at that time. Increased albedo
values over sub-tropical areas are important in
leading to the extension of arid conditions. In-
creasing the albedo of a surface contributes to
a net loss of radiative heat thereby creating
a horizontal atmospheric temperature gradient
with respect to the surrounding areas. Through
this a circulation is set up which transfers heat
aloft and maintains thermal equilibrium through
sinking and adiabatic compression (Charney
1975). In sub-tropical areas this subsidence is
superimposed on the descending branch of the
Hadley Cell, clearly accentuating its effect. The
positive feedback relationship which is set up as
the result of increased albedo values almost
implies that “the desert feeds upon itself”
(Charney 1975).

A number of other considerations also point
to the fact that during full glacial conditions
much of Western Australia must have been much
drier than today. Analyses of Indian Ocean cores
(Vella et al. 1975; Williams and Johnson 1975;
Be and Duplessy 1976) show that during the
last full glacial an equatorward shift of the Aus-
tralian-Subantarctic front and the Sub-Tropical
Convergence zone took place, with the result that
much lower sea surface temperatures character-
ised the ocean off Western Australia. Microfaunal
and oxygen isotope analysis of core RC9-150 (Be
and Duplessy 1976), taken from the continental
slope 125 km north-west of Perth, showed that
mean summer sea surface temperatures at 18 000
year BP were up to 6C° lower than those of
today.

Sea surface temperatures exert an important
control on precipitation (e.g. Namias 1975;
Wright 1977), and in tropical areas show a
strong correlation with storm frequencies
(Wendland 1977). Today the presence of warm
water off the coast in winter makes south-west-
ern Australia far wetter than any correspond-
ing western littoral (Gentilli 1952, 1971; Lydolph
1957). An examination of the rainfall distribu-
tion of the dry western littorals of the various
continents (Lydolph 1957), revealed that except

138

Journal of the Royal Society of Western Australia, Vol. 62, Parts 1-4, 1979.

for Western Australia the annual rainfall
increases inland from the driest area which lies
in a narrow coastal strip. This extreme aridity
in the immediate coastal belt may be explained
by the stabilising effect of a cold offshore current,
which today is largely absent off the coast of
Western Australia (Wooster and Reid 1963). In
the Indian Ocean the low-latitude ocean circula-
tion is strongly determined by the trade winds
(Knauss 1963). With an increase in the strength
of the trades during full glacial periods, which
now seems likely in the light of evidence from
different parts of the world (e.g. Parkin 1974;
Molina-Cruz 1977) a cold offshore current along
the coast of Western Australia can be expected.
In any event with the considerably lower sea
surface temperatures, that the deep-sea core
evidence suggests, a tendency towards aridity
should have been pronounced. This conclusion
was in part anticipated by Brooks (in Mayr 1944)
over 30 years ago. He suggested that during full
glacial conditions a cold current was present off
the west coast of Australia, which he suggested
would only be a winter-time phenomenon. The
presence of this current “would have had the
effect of extending the winter-time high pressure
area and contributing at this season to the
aridity of the west coast of Australia” (Mayr
1944, p. 129).

The existence of arid conditions over parts of
Western Australia during full glacial periods
would have been further facilitated by the ex-
posed continental shelves of those periods. That
the increased continentality would result in an
appreciable reduction in the precipitation re-
ceived by the north-west was again pointed out
by Brooks, and more recently reiterated by Nix
and Kalma (1972). The important control that
this mechanism is likely to have had on the late
Quaternary climates of the Nullarbor Plain
(Martin 1973; Martin and Peterson 1978) was
noted in an earlier section.

Summary

Despite the sketchy nature of the evidence it
is now apparent that Western Australia experi-
enced widespread climatic changes over the last
80 000 years. Of these changes the most severe
was clearly the arid phase which characterised
part of the closing 10 000 years of the Pleistocene.
Arid-zone extension, coinciding with the global
glacial maxima, must have been a recurrent
feature of the Pleistocene climatic history of
Western Australia. This inference is in part
supported by the stratigraphy of the Quaternary
sediments of the Shark Bay-Exmouth Gulf area.
But it needs to be borne in mind that aridity in
Australia finds its origin in the late Tertiary
(see Bowler 1976), so that the Pleistocene exten-
sions of the arid zone are the culmination of a
trend which started some 10 million years ago.
The extreme arid phases of the Pleistocene must
have deeply influenced the lives of the original
human inhabitants of this part of the continent,
and their effect on both flora and fauna was
equally profound. Similarly much of the physi-
cal landscape still bears the imprint of these
phases.

Acknowledgements. — I am grateful to W. J. E. van de
Graaff, D. Merrilees, J. Gentilli and I. Watson for con-
structive criticism of an earlier draft of the manu-
scrip . A. B. Pittock was most helpful in commenting
on the use of the “Z” criterion. I thank D. Glassford
for discussion. Without the help of my wife this paper
would not have been completed.

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Appendix 1

The partial equivalent potential temperature
is given by

0 PE = Kd EA1]

where e is the potential temperature. If air at
pressure p and temperature T is brought adia-
batically to a standard pressure P<> of 1 000 mb.,
its temperature e at that standard pressure is
known as its potential temperature. This is
given by

e = T (Po/P ) K t A2]

K = (.Cp-Cv) / cp = 0.288 for dry air
cp is the specific heat at constant pressure
cv is the specific heat at constant volume

In equation [All the function K introduces
the modification in the potential temperature due
to the presence of moisture in the air; for
details of this see Greenhut (1977).

142

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Journal

of the

Royal Society of Western Australia
Volume 62 1979

Parts 1-4

Contents

Page

History of geology in Western Australia. By J. H. Lord 3

Stromatolite research in Western Australia. By P. E. Playford 13

The contribution of geochronology to Precambrian studies in Western

Australia. By J. R. de Laeter and A. F. Trendall 21

History of zoology in Western Australia. By D. L. Serventy 33

The Western Rock Lobster, Panulirus cygnus. By R. W. George, G. R.

Morgan and B. F. Phillips 45

The breeding seasons of birds in south-western Australia. By S. J. J.

F. Davies 53

Niche theory and macropodid nutrition. By J. E. Kinnear and A. R.

Main 55

Vegetation mapping in Western Australia. By J. S. Beard 75

Plant physiologist amongst the grain legumes. By J. S. Pate 83

Studies on the resin chemistry of some Western Australian plants. By

P. R. Jefferies 95

The prehistoric environment in Western Australia. By D. Merrilees 109

Late Quaternary climates of Western Australia: evidence and

mechanisms. By K-H Wyrwoll 129

Editor: A. E. Cockbain

The Royal Society of Western Australia, Western Australian Museum, Perth

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