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THE

PIO IE JE DVO Ges

OF THE

L_INNEAN SOCIETY

OF

NEW S@Urom VV ALES

FOR THE YEAR

19383

VOL. LVIII

WITH TWENTY-NINE PLATES
and 319 Text-figures.

SYDNEY:
PRINTED AND PUBLISHED FOR THE SOCIETY BY

AUSTRALASIAN MEDICAL PUBLISHING CO., LTD,
Seamer Street, Glebe, Sydney,

and
SOLD BY THE SOCIETY.
1933.

ii CONTENTS.

CONTENTS OF PROCEEDINGS, 1933.

PARTS I-II (Nos. 245-246).
(Issued 15th May, 1938.)

Presidential Address, delivered at the Fifty-eighth Annual Meeting,
29th March, 19338, by C. Anderson, M.A., D.Sc. ..

Elections
Balance-sheets for the Fourteen Months ending 28th February, 1933 ..

The Mayflies of the Mount Kosciusko Region. i. (Plectoptera.)
Introduction and Family Siphlonuridae. By R. J. Tillyard,
M.A., Se.D., D.Se., F.R.S. (Plate i and forty-five Text-figures.)

The Life-history of Grevillea robusta (Cunn.). By P. Brough, M.A.,
B.Sc., B.Sec.Agr. (Plate ii and ninety Text-figures.)

Notes on Australian Diptera, XXxXiii. By J. R. Malloch.
(Communicated by F. H. Taylor.) (One Text-figure.)

Revision of Australian Lepidoptera. Oecophoridae. ii. By A. Jefferis
Turner, M.D., F.E.S.

On the Production of Fertile Hybrids from Crosses between Vulgare
and Khapli Emmer Wheats. By W. lL. Waterhouse, D.Sc.Agr.
(Plates iii-iv.)

Some Climatological Aspects of Aridity in their Application to
Australia. By John Andrews, B.A., and W. H. Maze. (Seven
Text-figures.) ..

Seasonal Incidence and Concentration of Rainfall in Australia. By
John Andrews, B.A., and W. H. Maze. (Five Text-figures.) ..

Pages.

1—-XXvV
XXV

XXVI-XXVili

1— 32
ae (8!
74— 79
80- 98
99-104

105-120
121-124

CONTENTS.

PARTS III-IV (Nos. 247-248).

([ssued 15th September, 1933.

The Petrology of the Hartley District. ii, The Metamorphosed Gabbros
and Associated Hybrid and Contaminated Rocks. By Germaine A.
Joplin, B.Sc. (Plates v—vi and four Text-figures. )

Australian Coleoptera. Notes and New Species. viii. By H. J. Carter,
B.A., F.E.S. (Hight Text-figures. )

Corynebacteria as an Important Group of Soil Microorganisms. By H. L.
Jensen, Macleay Bacteriologist to the Society ..

The Marine Plankton of the Coastal Waters of New South Wales. i. The
Chief Planktonic Forms and their Seasonal Distribution. By
Professor W. J. Dakin, D.Sc., F.Z.S., and Allen Colefax, B.Sc. (Plate

vii and seven Text-figures.)

Notes on New South Wales and Queensland Orchids. By Rev. H. M. R.
Rupp, B.A. (Two Text-figures.)

The Surface History of Monaro, N.S.W. By Frank A. Craft, B.Sc.,
Linnean Macleay Fellow of the Society in Geography. (Plate ix and
four Text-figures. )

Vegetative Reproduction in Drosera peltata and D. auriculata. By
Joyce W. Vickery, M.Sc. (Plate viii and thirty-four Text-figures. )

On the Distribution, Habitat and Reproductive Habits of Certain
European and Australian Snakes and Lizards, with Particular Regard
to their Adoption of Viviparity. By H. Claire Weekes, D.Sc.,
Linnean Macleay Fellow of the Society in Zoology. (One Text-
figure.)

A New Genus and Species of Australian Proctotrypidae. By Alan P. Dodd.
(One Text-figure.)

The Scientific Name of the Commercial Oyster of New South Wales. By
Tom Iredale and T. C. Roughley ..

The Life History of the Australian Oyster (Ostrea commercials). By
T. C. Roughley, B.Se., F.R.Z.S. (Plates x-xxvii and two Text-
figures. )

iii

Pages.

125-158

159-180

181-185

186-222

223-228

245-269

279-333

iv CONTENTS.

PARTS V-VI (Nos. 249-250).
(Issued 15th December, 1933.)

Pages.
The Geology of the South Coast of New South Wales, with special
reference to the Origin and Relationships of the Igneous Rocks. By

Ida A. Brown, D.Sc., Linnean Macleay Fellow of the Society in

Geology. (Plate xxviii and four Text-figures.) .. .. 3884-362
The Coccidae of the Casuarinas. By Walter W. Froggatt, F.L.S. (Plate

xxix and thirty-six Text-figures. ) Go naan ete ena aa) Oca == SSI
An Investigation of the Sooty Moulds of New South Wales. i. Historical

and Introductory Account. By Lilian Fraser, M.Sc., Linnean Macleay

Fellow of the Society in Botany .. ma : AY ; 375-395
Notes on the Australian Species of the Family Paussidae (Coleoptera).

By the late T. G. Sloane (Prepared for publication by H. J. Carter,

B.A., F.E.S.) ity atauie Maes (badge TONES se cole ee ail ee eae ee OO OOS
Useful Coccinellidae found on the Comboyne Plateau. By E. C. Chisholm,

M.B., Ch.M. (Hight Text-figures.) APD scree dee aca: ke ed FAO iT,
Miscellaneous Notes on Australian Diptera. i. By G. H. Hardy. (Four

Text-figures. ) 408-420
The Genus Pterostylis R.Br. (oid A new Scheme of Classifica-

tion, with Notes on the Distribution of the Australian Species. By

the Rev. H. M. R. Rupp, B.A. (Thirty-eight Text-figures.) 421-428
A new Species of Pterostylis. By (Mrs.) Pearl R. Messmer. (Ten Text-

figures. ) dee 1 Thee BR Gp ea She a Rene tte ee RRA = Ae} ()
The Mycetozoa of New South Wales. By Lilian Fraser, M.Sc., Linnean

Macleay Fellow of the Society in Botany Ae EE Rt eee A SAS 6
The Coastal Tablelands and Streams of New South Wales. By Frank A.

Craft, B.Se., Linnean Macleay Fellow of the Society in Geography.

(Six Text-figures.) Sibiu Fiat VSG . .. 437-460
Australian Hesperiidae. iv. Notes and Descriptions of new Forms. By

G. A. Waterhouse, D.Sc., B.E., F.E.S. PUAN cual) ae Warn tele Wad O46 G
Some Further Observations on the Rearing of Ceratodus. By Thos. L.

Bancroft, M.B., Ch.M., C.M.Z.S. (Communicated by I. M. Mackerras,

M.B., B.Sc.) (Two Text-figures.) 467-469
Abstract “of MP roceedin gsm yee nn) mee! ee oy esis inlet WI Paha il eee yale XXiX—xXXXVi
Donations and Exchanges pen ee aM re Re UU Pee IRR en Mate Bs led O.0, QVC >.dU.<
List of Members l-liv
Index ; lv—-lxiii
List of New Genera .. lxiii
Corrigenda lxiv

List of Plates

lxiv

ANNUAL GENERAL MEETING.
WEDNESDAY, 29th Marcu, 1933.

The Fifty-eighth Annual General Meeting was held in the Society’s Rooms,
Science House, Gloucester Street, Sydney, on Wednesday, 29th March, 1933.

Dr. C. Anderson, M.A., President, in the chair.

The minutes of the preceding Annual General Meeting (30th March, 1932)
were read and confirmed.

PRESIDENTIAL ADDRESS.
Ladies and Gentlemen,

I have the honour to address you on the occasion of our Fifty-eighth Annual
Meeting, and it is my pleasure to assure you that in spite of the prevailing
depression, which has affected our finances to a certain extent, there has been
no slackening in the successful carrying on of our appointed work.

The concluding Part of Volume lvii of the Society’s ProcrEpIncs was issued
in December, in accordance with the Council’s decision to publish the year’s
proceedings in three issues instead of six as formerly. The complete volume (410
plus lxvii pages, nine plates and 227 text-figures) contains thirty-six papers from
twenty-nine authors, six papers being by Linnean Macleay Fellows and two by
the Macleay Bacteriologist. The reduced size of the volume was a result of the
difficult financial year experienced by the Society. It is hoped that conditions
will show some improvement during 1933, but authors would still assist the
Council considerably by continuing to condense their papers into the smallest
space.

Exchanges from scientific societies and institutions were well maintained
and receipts amounted to 2,236 for the session, compared with 2,084, 1,866 and 1914
for preceding years.

During the year Special Meetings were held to alter the Rules, so that the
Society’s financial year would commence on the first day of March in each year
instead of the first day of January. This makes the Society’s financial year
correspond with the financial year of Science House and also with the year of the
Linnean Macleay Fellowships.

At the same time the rule governing Corresponding Membership was altered
so as to empower the Council to elect persons resident in New South Wales
as Corresponding Members. Following this alteration the Council elected Mr.
W.S. Dun, a past President of the Society, as a Corresponding Member.

The twenty-first meeting of the Australian and New Zealand Association for
the Advancement of Science was held in Sydney in August, 1932, under the
Presidency of His Excellency Sir Hubert Murray, Lieutenant-Governor of Papua.
As the Association had not met in Sydney since 1911, the meeting was expected
to be a very successful one, and expectations were amply fulfilled, the membership
for the meeting reaching a total of 1,085, a very satisfactory result for these
difficult times. It is indeed gratifying to find such a measure of support for
science under the prevailing conditions of financial stringency.

A

ii PRESIDENTIAL ADDRESS.

As one result of explorations in which this Society has always taken an
interest on account of the participation of several of its members, a large section
of the Antarctic continent has been placed under the authority of the Common-
wealth by a British Order-in-Council. The Australian claim to this responsibility
was based largely on the rights of discovery and exploration. Sir Douglas
Mawson’s expeditions of 1911-14 and 1929-31 had explored portions of the
continent previously known and had also added important new discoveries. The
sector now placed under Australian control comprises the lands of the Antarctic
Continent between longitudes 45° EH. and 160° E. with the exception of Adélie
Land. In this region Mawson’s expeditions visited King George V Land, Wilkes
Land, Queen Mary Land, Enderby Land, Kemp Land, and the recently discovered
and named lands, MacRobertson Land and Princess Elizabeth Land. In this
new responsibility there are commercial possibilities in the development of the
whaling industry and the fisheries.

An unusual compliment was paid to one of my predecessors in this Chair,
Dr. H. S. H. Wardlaw, in a request that came last year from the Smithsonian
Institution for permission to reprint in its Annual Report his Presidential Address
to this Society.

The recent appointment of Mr. EK. Cheel as a member of the National Park
Trust will give satisfaction to those interested in natural history. There is ample
need for a small proportion of representatives of science on such a Trust which
has to administer a large area which is a National Reserve where every effort
should be made to preserve the native fauna and flora. Mr. Cheel has for years
taken a keen interest in the flora of National Park, to the study of which he has
devoted much of his leisure time, and this interest in the Park may now prove
to be of some practical use to the Trust.

During the past year the Council appointed a small standing Committee,
consisting of Messrs. Cheel, Hamilton and Froggatt, to consider what might be
done to popularize our native flora.

A year ago your President reported that a deputation to the Prime Minister
had been unsuccessful in its efforts to secure the removal of the Sales Tax from
books of an educational character. It is pleasing to be able now to report that
the Federal Government took the opportunity in November last of placing
educational books on the list of exemptions and thus removing what had proved
a serious handicap to scientific societies and institutions in maintaining their
libraries.

The Catalogue of Scientific and Technical Periodicals published in 1930 by
the Council for Scientific and Industrial Research has proved so useful that
suggestions have been made in many quarters for the issue, from time to time,
of supplements. Following a grant made by the Australian and New Zealand
Association for the Advancement of Science towards the publication of such a
supplement, scientific societies and institutions were invited by the Council for
Scientific and Industrial Research to contribute towards the cost of the publication,
and it is satisfactory to know that sufficient support was forthcoming for the
work to be put in hand.

In August last a conference took place in Sydney at which representatives
of the various Royal Societies in Australia and of this Society were present,
the object being to discuss the desirability of some sort of affiliation between
the Societies and the Australian National Research Council. After considerable
discussion the delegates unanimously carried the following resolution: “That

PRESIDENTIAL ADDRESS. iii

the delegates to this conference are of the opinion that an amalgamation of the
Societies represented is not desirable, but they recommend that some kind of
loose federation be formed such as could be obtained by each Society nominating
a member to the executive of the Australian National Research Council.”

Since the last Annual Meeting nineteen Ordinary Members have been added
to the roll, five have resigned, the names of three have been removed on account
of arrears of subscription, and two have been lost by death. One Corresponding
Member has died during the year.

THOMAS STORIE Dixson, who died on 9th December, 1932, was born in 1854
and was a son of Hugh Dixson, founder of the firm of Dixson & Sons, tobacco
manufacturers. At the age of eighteen he commenced the study of medicine at
the University of Edinburgh, taking the degree of M.B. in 1877. He intended
also to graduate in Science, but ill-health prevented this. He studied widely
abroad, at Dublin, Berlin, Vienna and Strassburg, and translated into English
Schmiedeberg’s “Pharmacologie”’ which became a standard text-book in English-
speaking medical schools. He was for a number of years after his return to
Sydney Lecturer in Materia Medica and Therapeutics at the University of Sydney.
He was one of the best known medical men in New South Wales, having, during a
period of nearly fifty years, filled many public positions and rendered wonderful
service to many charitable institutions in Sydney. Dr. Dixson was President
of the Medical Board from 1919 till the time of his death. Though he did not
graduate in Science he retained his interest in natural history, as evidenced
by his connection with the Australian Museum and with this Society. He was a
member of the Board of Trustees of the Museum from 1898 till his death, and
its President from 1918 to 1925. He became a member of this Society in 1881,
and was elected a member of Council in 1882. He was President for the years
1903-4 and 1904-5, and Vice-President for the years 1902-3 and 1905-11. He was
also one of the Honorary Secretaries for the year 1892-3. He made only one
contribution to the ProcrEpInes (1882) in addition to his Presidential Addresses.
His period of fifty years’ service as a member of Council constitutes a record
which may stand for a long time. That he sustained his interest and regularly
attended meetings of the Council, except during his last year or two, was no
doubt due to the fact that he was a personal friend of our great benefactor,
Sir William Macleay. His death removes one of the very few members left
to us who knew William Macleay personally.

THoMAS GIBSON SLOANE, who died at the Burrangong Hospital, Young, on
20th October, 1932, was born in Melbourne on 30th April, 1858. He was one of five
sons of Alexander Sloane of Mulwala, Murray River, and was educated partly in
Melbourne at Scotch College and partly at home by tutor. Sloane followed in the
footsteps of his father, who was a prominent merino expert, and in 1888 became
manager of Moorilla Station. He took the keenest interest in sheep-breeding, won
numerous prizes at sheep shows, and kept methodical records of his wool weights.
He had a stud of his own registered in the Flock Book. Prior to taking over
management of Moorilla he visited Sydney to learn business methods and came
to know, through his interest in natural history, Sir William Macleay and J. J.
Fletcher, forming a lifelong friendship with the latter. As an entomologist he
was recognized as the authority in Australia on the Carabidae and Cicindelidae,
though he did not confine his systematic work to these two groups. He had too
wide an interest, however, to be content with descriptive entomology, and
took great interest in problems of phylogeny and distribution, some of his papers

iv PRESIDENTIAL ADDRESS.

being notable additions to the literature of these subjects, particularly ““‘The Faunal
Sub-Regions of Australia’ (1915) and “Classification of the Family Carabidae”
(1923). He published sixty-one papers, the first appearing in 1881 and the
last in 1928. Of these, forty-nine were published in our PRocEEDINGS in the years
1888-1923. He himself collected widely, not only in New South Wales and Victoria,
but also on extended trips to Western Australia and Queensland, and as a result
his collection was an extensive one, containing the types of many of the species
he described as new. His collection has been presented by Mrs. Sloane to the
Council for Scientific and Industrial Research, and will be preserved at Canberra.

He was one of the oldest members of the Society, having joined in 1887,
and though we did not often see him at meetings, it was his habit to visit the
Society’s rooms on his periodic visits to Sydney for the sheep shows up till 1924,
when he practically gave up his scientific work. No doubt this habit was formed
during his lifelong friendship with J. J. Fletcher.

DANIEL McALPINE, a Corresponding Member of the Society since 1902, who
died at Leitchville, Victoria, in October, 1932, was born on 21st January, 1849:
He came to Australia in 1884 and was appointed Lecturer in Biology at Ormond
College, Melbourne University, and Lecturer in Botany at the College of Pharmacy,
Melbourne. In 1890 he became Plant Pathologist to the Victorian Government,
and in 1911 was appointed as Commonwealth Commissioner to investigate the
nature and control of bitter pit in apples. He retired in 1915. He attained a
very wide reputation for his work in plant pathology and at the Pan-Pacific
Congress held in Australia in 1923 a resolution was passed expressing regret
that he was unable to attend and also expressing deep appreciation of the value
of his contributions to plant pathology. His publications were exceedingly
numerous, totalling well over two hundred, of which perhaps his monographs
on the Rusts and Smuts of Australia and on Bitter Pit may be singled out for
special reference.

I may be permitted to refer also to the death of one who was an active
member of the Society for many years.

Dr. N. A. Coss, Principal Nematologist to the Bureau of Plant Industry,
U.S. Department of Agriculture, died on 4th June, 1932, having almost completed
his 73rd year. He was born in Spencer, Mass., on 30th June, 1859. He received
his doctor’s degree at Jena in 1888 and after spending a season at the Zoological
Station, Naples, came to Australia early in 1889. He was appointed to act in
Professor Haswell’s place during the latter’s absence on leave, and in 1891
became pathologist in the New South Wales Department of Agriculture, where he
remained for eleven years. In 1898-1900 he went abroad to the United States of
America, Europe and Algeria, having been appointed a Special Commissioner to
report on Agriculture in America and Europe. He left Australia in 1905 and
became director of Physiology and Pathology to the Hawaiian Sugar Planters’
Experiment Station and in 1907 was appointed Agricultural Technologist in
the United States Department of Agriculture. He was a member of this Society,
1889-1906, and contributed eight papers, chiefly on Nematodes, to the Proceedings
during the years 1890-1898. He was also a member of Council during the years
1892-1894.

The publication during the past year of several books by members of the
Society seems worthy of reference on this occasion. ;

Sir Edgeworth David’s Geological Map of Australia, accompanied by a
volume of explanatory notes, published by the Council for Scientific and Industrial

PRESIDENTIAL ADDRESS. v

Research, marks a definite step towards the publication of his book on the Geology
of the Commonwealth, which is approaching completion. The preparation of the
map has involved a tremendous amount of work which no one has been better
fitted to do than Sir Edgeworth, while the accompanying volume forms an
admirable summary of the Geology of Australia.

Dr. G. A. Waterhouse’s “What Butterfly is That?” (published by Messrs.
Angus & Robertson, Ltd.) demonstrates again the ability of Australian scientist,
artist and printer to combine and produce an adequate and well illustrated volume
on some phase of Australian Natural History. By the careful and methodical
descriptions of the species, and of their life-histories where known, and the
excellent coloured reproductions of the species the author surely has attained his
object of enabling ‘‘anyone to identify readily any butterfly he is likely to see
in the settled parts of Australia’. And his interesting peeps into the life-
histories should just as surely be the means of encouraging students to add
to the sum of our knowledge of the life-histories of butterfly species, and to help
complete our knowledge of their distribution. Dr. Waterhouse is to be con-
gratulated on the production of a volume which, whilst avoiding the unnecessary
use of technical terms, will be of great value to those who have more than a
superficial knowledge of our butterfly fauna. The artists, Mrs. D. S. North and
Mr. Neville Cayley, deserve a word of praise for the excellence of their work.
The twenty-five coloured plates are on the whole excellently produced, but it
would appear that something is yet to be learned of the technique necessary to
attain exact registration of the colours over the whole area of the plate.

Mr. A. Musgrave’s “Bibliography of Australian Entomology, 1775-1930”,
published by the Royal Zoological Society of New South Wales, is one whose
value to scientific workers is inestimable. Whilst it is intended primarily for the
entomologist, it is certain that those interested in other branches of natural
history will find in its pages much that is of interest. The compilation of the
volume has entailed a vast amount of careful work which the author has per-
formed faithfully and well, and he will reap his reward in the knowledge that
not only the present, but future, generations will find many of their labours
lightened as a result of his work.

Mr. R. H. Anderson’s “The Trees of New South Wales”, published by the
Department of Agriculture, N.S.W., is intended to be of use to the many people
who are interested in our trees, but who may lack the technical knowledge of
such people as the botanist and the forester. Brief non-technical descriptions of
the trees are given, and there are illustrations of many of the commoner and
more interesting species.

The year’s work of the Society’s research staff may be summarized thus:

Mr. H. L. Jensen, Macleay Bacteriologist to the Society, completed the study
on the sixty-seven strains of Micromonospora. They were found to fall into four
groups, of which three represented new species, and were found to occur most
frequently in neutral to alkaline soils from districts with a low rainfall. The
results have been published as Part iii of the series of papers on the Actino-
mycetales. Periodical counts of numbers of bacteria, actinomycetes and fungi in
the soil have been carried out, together with similar counts in a number of
soil samples from various parts of New South Wales. The results, although not
yet complete, indicate that the balance between bacteria and actinomycetes is
governed chiefly by the moisture content of the soil, the actinomycetes becoming
relatively more numerous during dry periods. The actual numbers of all three

vi PRESIDENTIAL ADDRESS.

groups of organisms seem, under approximately uniform conditions of moisture,
to depend mostly on the supply of organic matter in soil and, so far as the
bacteria and actinomycetes are concerned, upon the reaction. In dry soils poor
in organic matter the numbers of fungi are strikingly low. A considerable amount
of work has been carried out on the mycobacteria and corynebacteria of the
soil. The latter group, the most well-known representative of which is the
diphtheria bacillus, has hardly been studied at all from other than medical points
of view. The results obtained here show that these organisms, so far from
representing a narrow and specialized group of bacteria parasitic in man and
warm-blooded animals, do really occur abundantly as saprophytes in nature and
particularly in soil. A preliminary paper has been completed in which it is
shown that corynebacteria account for a large proportion, even up to two-thirds,
of the soil bacteria which develop on agar plates. They are doubtless identical
with organisms sometimes mistaken for nodule bacteria of leguminous plants.
Most of the mycobacteria found here correspond to previously known types,
although not all well described, but the collection of corynebacteria, which numbers
about forty strains, seems to represent several new species. Some types of
both genera represent a very close transition to the genus Proactinomyces and
produce through phenomena of dissociation, i.e., mutation-like changes, variants
indistinguishable from this genus. Such phenomena may both occur spontaneously
and be induced experimentally. Several cultures of alleged mycobacteria and
related organisms have been obtained for comparison with the forms isolated
here, and some of them have been found to have their proper place in the genus
Proactinomyces. A paper on the identity of these organisms has been published
as Part iv of the series of papers on the Actinomycetales.

Dr. Ida A. Brown, Linnean Macleay Fellow of the Society in Geology, carried
out further extensive field work in the South Coast district. In the older Palaeozoic
(pre-Devonian) rocks between Quaama, Bermagui, Narooma and Bateman’s Bay
she found two distinct series of rocks, the one consisting of quartzites and
graptolite-bearing slates, definitely Upper Ordovician in age, and the other of meta-
morphosed sediments and igneous rocks bearing striking resemblance to the
Brisbane “Schist’” Series of Queensland. Of the pre-Devonian rocks from the
head of the Clyde River to the Victorian border (about 150 miles) and for 20
or 30 miles inland from the coast, only those in the vicinity of Bendithera are
of (Upper) Silurian age; apart from the Upper Ordovician graptolite-bearing
rocks which outcrop over small areas near Cobargo and Quaama, the remainder
of the series consists of rocks which Dr. Brown believes to be older than Upper
Ordovician. They consist of black banded radiolarian cherts associated with
pillow lavas, volcanic agglomerates, etc., whose characters suggest correlation
with Cambrian formations in Victoria and possibly also with the Cobar-Canbelego
Series in New South Wales and portion of the Brisbane Schist Series in Queens-
land. She has in preparation a paper on these Older Palaeozoic rocks of the
South Coast which appear to have formed a “borderland” to the continental
massif during early Palaeozoic time, and the study of this problem has led her
to a reconsideration of the structure and tectonic history of South-eastern Australia
from Cambrian (?) to Upper Devonian time. One paper appeared in the
ProcEeepines for 1932, viz., “Late Middle Devonian Diastrophism in South-eastern
Australia’, and in this evidence is produced of important earth-movements in
South-eastern Australia between the deposition of Middle and Upper Devonian
sediments. The main results of Dr. Brown’s work as a Linnean Macleay Fellow

PRESIDENTIAL ADDRESS. vii

were collected in a thesis entitled ‘‘The Geology of the South Coast of New South
Wales, with Special Reference to the Origin and Relationships of the Igneous
Rocks”, which gained for her the degree of Doctor of Science from the University
of Sydney. This thesis, which has now been submitted for publication, deals
with problems of the geological age, conditions of sedimentation, mutual relation-
ships and subsequent tectonic history of the sedimentary rocks, and the relation-
ships, petrogenesis and correlation of the associated igneous rocks. She has
also two short papers in preparation, in completion of her work, dealing with
the Upper Devonian rocks of Mount Lambie, and the Tertiary and Post-Tertiary
Geology and Physiography of the South Coast. Dr. Brown retired from her
Fellowship on 28th February last; the results of her work during the six years
in which she was a Linnean Macleay Fellow form a very important addition
to our knowledge of the geology of the South Coast district of New South Wales
and, indeed, of the South-east of Australia. I may take this opportunity on behalf
of the Society of wishing her every success in the future.

Mr. F. A. Craft, Linnean Macleay Fellow of the Society in Geography, carried
out field work in the Monaro region, covering a large area of country which
included the district from Cooma to the head of the Murrumbidgee River, the
upper valley of the Hucumbene, part of the Snowy River drainage to the south
and south-west of Cooma, the tableland east of Cooma, and the routes between
Cooma and Goulburn. As a result of this he has completed a paper, “The
Surface History of Monaro, N.S.W.”, in which he discusses the typical land forms
and topography and their evolution, and also the development of the stream system
which he holds to be inseparable from that of Hastern Victoria. Then he carried
out a reconnaissance of southern New England, which enabled him to complete a
paper on “The Coastal Tablelands and Streams of New South Wales”, in which
most attention is devoted to the eastern slopes of the plateau and the Pacific
streams, but some reference is made to the western slopes and features. During
1932 four papers appeared in the Procrrpines, including the final two parts of
his series on ‘‘The Physiography of the Shoalhaven River Valley’, as well as
“Geographical Studies in the Blue Mountain Tableland” and ‘Notes on Erosional
Processes and Stream Gravels”’. During the coming year he proposes to study
erosional problems in the New South Wales part of the Murray River catchment
above the Hume Dam. This will include a consideration of the land forms of
the Upper Murray Basin in this State, the nature and extent of erosion in various
parts of the terrain, estimates of rates of erosion in various parts of the landscape
and the consequent modification of land forms and stream profiles, making allow-
ance for effects due to settlement. This should be a contribution to physiography,
both as regards erosion itself and the fate of the eroded material. Some work
has already been done by Mr. B. U. Byles, now of the State Forestry Service,
who carried out a six months’ reconnaissance of the mountainous part of the
area and whose reports have been very generously handed over to Mr. Craft
for his use.

Dr. H. Claire Weekes, Linnean Macleay Fellow of the Soicety in Zoology,
commenced by experimenting with new technique for the physiological and
histological investigation of the pituitary, adrenal bodies, thyroid and ovary otf
lizards during hibernation and the various stages of the breeding season. She
spent considerable time collecting material on the New England Plateau, North
Coast, Bathurst Plains, Blue Mountains, Coastal Districts round Sydney, South
Coast and Kosciusko. During these trips she collected lizards in all stages oz

viii PRESIDENTIAL ADDRESS.

adaptation to viviparity—lizards which lay eggs, lizards which are viviparous but
which have no reduction in the yolk-content of their eggs and only very simple
placentation, lizards which are viviparous and have a reduction in the yolk-
content of their eggs and a comparatively well specialized placenta. Microtome
sections were prepared of the material collected, and during February observa-
tions were made on pregnant lizards and the birth of the young. She has
completed for publication a short paper on the evolution of viviparity among
certain reptiles, and also has ready a paper giving a survey of placentation among
reptiles and its evolution, and possible causes of viviparity—portion of which
was read before the Anatomical Society of Great Britain and Ireland in London
in 1931. During the coming year she proposes to follow the work done on
placentation and viviparity by an investigation of the endocrine glands concerned
in the physiology of these specialized phases of the reproductive cycle. From
the material collected during 1932 she also hopes to make a comparative study
of the pituitary, adrenal bodies, thyroid and ovary of the lizards with the
object of discovering, if possible, the nature of the mechanism or mechanisms
controlling the retention of the eggs within the oviduct and the ejection of the
eggs or young at birth.

Miss Lilian Fraser, Linnean Macleay Fellow of the Society in Botany, has
been working on the sooty moulds of New South Wales. She has investigated
the types of fungi present in the various moulds encountered, and found that the
sooty mould (as in the Northern Hemisphere) consists of a number of fungi
growing in conjunction to form a community. In general, the associated fungi
were found to be similar to those found elsewhere, but there was a notable
absence of some of the common moulds. Monthly examinations were made of
sooty moulds growing on selected trees in order to find out whether there are
seasonal changes in the composition of the mould, the results so far indicating
the presence of more types of fungi during the wet autumn and winter months.
Experiments have been made on a selected number of fungi, grown in culture,
to investigate their food requirements. A number of fungi have been used and
they have been grown in a variety of culture media. These experiments have
been completed, but the results have not yet been fully analysed. Experiments
are being carried out to find the effect of direct sunlight on the growth of the
fungi used in the growth studies, together with a number of others obtained in
culture from sooty moulds. Attempts made to obtain germination of seeds of
Lobelia gibbosa and L. dentata have so far not been successful. During 1932 she
completed, in conjunction with Miss Gladys Carey, a paper on “‘The Embryology
and Seedling Development of Aegiceras majus Gaertn.” which was published in the
PROCEEDINGS. She proposes to complete the physiological studies on the group,
and to make an examination of the life-history of Capnodium sp., and also to
investigate the biology. of a group of fungi collected from members of the
Proteaceae, particularly their distribution, their differential resistance to the
various genera and species of Proteaceae, methods of infection and seasonal
changes.

Five applications for Linnean Macleay Fellowships, 1933-34, were received
in response to the Council’s invitation of 28th September, 1932. I have pleasure
in reminding you that the Council reappointed Mr. F. A. Craft, Dr. H. Claire
Weekes and Miss Lilian Fraser to Fellowships in Geography, Zoology and Botany
respectively, and also appointed Dr. Ivor Vickery Newman, M.Sc., to a Linnean

PRESIDENTIAL ADDRESS. ix

Macleay Fellowship in Botany for one year from 1st March, 1933. I am sure you
will join me in wishing them a successful year’s research.

Dr. Ivor Vickery Newman graduated in Science at the University of Sydney
in 1926 with second class Honours in Botany. He then held a Government
Science Research Scholarship in Botany in 1927 and 1928, obtaining the degree
of M.Sc. as a result of research carried out during this period. He proceeded
to King’s College, University of London, in 1929, there working under Professor
R. R. Gates and obtaining his Ph.D. in 1931. His published work includes “The
Life-history of Doryanthes excelsa, Parts i and ii”, published in our PRrockeEpDINGS
for 1928 and 1929 respectively, and “Studies in Australian Acacias”, i (General
Introduction) and ii (The Life-history of Acacia Baileyana) recently published
in the Journal of the Linnean Society of London. His chief work for his year’s
Fellowship is to be a detailed examination of the cytology of Acacia Baileyana
from the point of view of (1) laying a foundation of chromosome morphology for
the study of evolutionary relationships in the genus, and (2) studying certain
aspects of cytological theory. He has in hand also material for a morphological
and cytological survey of species related to A. Baileyana.

Tue Fossir MAMMALS oF AUSTRALIA.

Introduction.

In the second portion of my address I propose to give a short account of
the first discovery of fossil mammals in Australia, which took place just about
one hundred years ago, and of the gradual accumulation of our present-day
knowledge of these interesting animals. I cannot hope to give an exhaustive
account in the space of this address, but it seems to me that the early history
of these discoveries is full of interest, and that I shall be doing some service
by recalling some of the salient features of the pioneering work of Ranken,
Mitchell, Leichhardt, Owen, Krefft, McCoy, and others, and of the subsequent
researches of more recent workers such as Etheridge, De Vis, Dun, Broom,
Chapman, Longman, Scott, Lord and Glauert, confining myself to the more
important points and indicating the main sources where fuller information may
be obtained.

Marsupials constitute the bulk of the described Australian fossil mammals,
but I make no attempt to enumerate all the genera and species that have been
recognized. This is rendered unnecessary by the recent publication of Dr. G. G._
Simpson’s exceedingly useful “Post-Mesozoic Marsupialia” (Simpson, 1930), which
contains a complete list and an extensive bibliography.

The earliest discoveries of fossil mammals in Australia have a_ special
importance in view of the peculiar features of the living fauna of the country,
and evidently they created a good deal of interest in scientific circles in Hurope.
A hundred years ago scientific thought as regards the evolution and succession
of life on the earth was in an interesting stage. The influence of Cuvier, the
founder of the comparative anatomy and palaeontology of vertebrates, who died
on 13th May, 1832, was still dominant, and his views as regards the geological
succession of life were widely accepted. It had been fully established that in
the geological past animals existed that are now extinct, that great faunal
changes had taken place from time to time, that whole faunas had disappeared
and been succeeded by others, and some reasonable explanation of this was
being sought. In explanation of the modifications that have taken place in
the character of animal life throughout the ages, Cuvier supposed that the earth

B

as PRESIDENTIAL ADDRESS.

had suffered a series of catastrophic changes in prehistoric times, as a result
of which the animals then living in a given country were destroyed, to be
succeeded by a different assemblage. He did not hold that life was created
anew after each convulsion, but that isolated portions of the earth escaped
the general destruction, and that from these centres the earth was populated
afresh. The latest of these catastrophes was, of course, the Noachian deluge.

A hundred years ago, Darwin, on board the Beagle on its memorable voyage
round the world, was making a series of observations which pointed to a different
explanation from that advanced by Cuvier, and indeed he was laying the founda-
tions for his great work on the origin of species which, however, was not published
until many years later. In South America Darwin was struck by the abundance
of Pleistocene mammals, and with the fact that, while these differed from the
living forms and were in part of gigantic dimensions, they were yet closely
related to the animals which still exist in South America. It was difficult to
reconcile this observation with Cuvier’s hypothesis of catastrophic destruction,
and to explain why there should be “this wonderful relationship in the same
continent between the living and the dead” (Darwin, 1886, p. 310). On the
other hand, on the theory of descent with modification, the succession of the
same types within the same area is at once explained in a quite natural manner.

The living mammalian fauna of Australia is even more distinctive than
that of South America, and it therefore became a matter of importance to
discover the nature of the mammals which were living here in the late Tertiary
and Pleistocene.

Early History.

The first discovery of fossil mammals in Australia was made at the Wellington
Caves, New South Wales, in or before 1830, and the discoverer seems to have
been George Ranken, of Kelloshiel, Bathurst, who is described in the quaint
language of the day as “that very respectable Colonist and Magistrate” (Lang,
1831, p. 365). Ranken explored the caves, descending by a rope which he had
fastened to what he thought was a projecting piece of rock, which, however,
broke and disclosed itself as a large bone. Other bones were found on the
floor of the caves and embedded in a kind of red clay which filled cracks and
openings in the limestone, the well-known bone breccia of the Wellington Caves.
Ranken made a small collection of bones and fragments and brought them to
Sydney with a view to their being sent to Edinburgh for examination by Professor
Robert Jameson, who then enjoyed a high reputation as a zoologist and geologist.

We can well appreciate the importance of a determination of the kinds of
animals represented by these bones. Were they identical with forms still
extant in Australia or were they of different genera and species? Were they
marsupials like the great majority of the existing Australian mammals, or were
they similar to the extinct forms described by Cuvier, Buckland, and others
from the Paris Basin, the Huropean caves, and elsewhere? These may seem
superfluous and trivial questions in the light of our present-day knowledge, but
at the time of which we are speaking Australia was almost a terra incognita,
and we can well understand the eager curiosity with which these old bones
were examined by zoologists and palaeontologists a hundred years ago.

In Sydney the bones came under the notice of the Rev. John Dunmore Lang,
one of the stormy petrels of Australian history, and the first printed account
of them appeared in the Sydney Gazette as an anonymous letter signed “L’”

PRESIDENTIAL ADDRESS. xi

and dated 21st May, 1830. This was subsequently printed in the Edinburgh New
Philosophical Journal as a communication by Dr. Lang. The author, while
disclaiming any authority as a comparative anatomist, concluded that the greater
number of the bones were not those of animals belonging to species now living
in Australia, and that they had been brought into the caves by some beast of
prey. He goes on to say that “while this very interesting discovery supplies us,
therefore, with another convincing proof of the reality and the universality of
the deluge, it supplies us also with a powerful motive of gratitude to Divine
Providence for that long-forgotten visitation. For if the territory were over-
run with such beasts of prey as the antediluvian inhabitants of the cave at
Wellington Valley it would not have been so eligible a place for the residence
of man as it actually is. The tiger or hyaena would have been a much more
formidable enemy to the Bathurst settler than the despicable native dog, though
indeed they would certainly have afforded a much nobler game to the gentlemen
of the Bathurst Hunt” (Lang, 1831, p. 368).

This collection of bones was duly forwarded to Jameson at Edinburgh, where
they were examined by himself and Dr. Adam, who found that “some of the
teeth were those of the wombat, some belonged to the kangaroo, others we
could not refer, from want of means of comparison. One bone, from its great
size, particularly arrested our attention, from its appearing to belong to an
animal larger than any of the living species in the Australian world. It appeared,
on comparing it with the splendid skeleton of the hippopotamus in the Museum,
to resemble the radial bone of that animal” (Jameson, 1831, p. 393). Jameson
submitted the collection to W. Clift, of the College of Surgeons, London, who
identified bones of the wombat, kangaroo, and dasyure, remarking that the
large bone bore a great resemblance to the radius of the hippopotamus and did
not belong to the elephant (Clift, in Jameson, 1831, pp. 394-5). Jameson
concluded “That New Holland was, at a former period, distinguished from the
other parts of the world by the same peculiarities in the organization of its
animals, which so strikingly characterize it at the present day”. And “That
the large bone resembling the radial bone of the hippopotamus, shows that
Australia formerly possessed animals much larger than any of the present
existing species, equalling or even exceeding in magnitude the hippopotamus;
a fact of high importance, when we recollect that the quadruped population of
New Holland is at present but meagre, the largest species being the kangaroo”
(pp. 395-6). i

These conclusions are quite sound, for you will observe that the large bone
was not stated to be actually that of a hippopotamus but merely as having a
resemblance to the radius of that animal.

The next communication is by Major Thomas L. (afterwards Sir Thomas)
Mitchell, Surveyor-General of New South Wales, who contributed a paper, dated
Sydney, 14th October, 1830, to the Geological Society of London (Mitchell, 1831).
Mitchell, a man of great ability and versatility, keenly interested in scientific
matters, had in the meantime visited the caves, apparently in company with
George Ranken, with whom he frequently corresponded, and had formed a
collection of the bones. His paper contained a short account of the caves and
the contained bones, which were again submitted to Clift and found to be those
of the kangaroo, wombat, dasyure, koala and phalanger, the most abundant
being those of the kangaroo. Along with these were two bones “not agreeing
with those of any of the animals at present known to exist in New South Wales.

xii PRESIDENTIAL ADDRESS.

The first and larger is supposed to belong to the elephant; the second is also
obscure and imperfect but seems to be a part of one of the superior maxillary
bones of an animal resembling the dugong; it contains a portion of a straight
tusk pointing directly forward.”

Ranken’s original collection and one formed by Mitchell were forwarded
to Cuvier at Paris, and were examined by him and by W. Pentland, who was
apparently associated with Cuvier in his work. These were reported upon by
Pentland (Pentland, 1831, 1832, 1833), who, in general, confirmed Clift’s deter-
minations, but identified the large bone as that of a small elephant (an identifica-
tion in which Cuvier concurred), and could find no bone that resembled that
of a dugong. We now know, of course, that both these bones were those of
Diprotodon, one of the most striking of the Post-Tertiary marsupials of Australia,
and that the fossil mammals of this continent tell the same story as do those
of the Pleistocene of South America, namely, they were allied to the forms
which are still extant.

Darwin was in Australia in 1836, and, although in the section of his
Journal of Researches devoted to Australia he makes no reference to the extinct
animals, he was possibly aware of these discoveries, for he was in personal
communication with Mitchell. Certainly in an earlier part of the same work
(which was written after his return to England) he makes reference to the
extinct marsupials of Australia (Darwin, 1884, p. 173), and in his Origin of
Species he refers to the fossil mammals of Australia as evidencing the succes-
sion of the same types within the same area during the later Tertiary periods
(Darwin, 1886, pp. 310-811). We see, therefore, that these early discoveries
are of considerable scientific and historical importance.

The first systematic account of the Wellington fossils we owe to Richard
Owen, then a young man in the early thirties, who contributed a description
to Mitchell’s Three Hxpeditions into the Interior of Eastern Australia (Mitchell,
1838, pp. 359-366). In this short account, which is illustrated by a number
of plates from drawings made by Mitchell, who was no mean artist, were estab-
lished a number of extinct species, including Diprotodon optatuwm, Macropus
(= Sthenurus) atlas, Macropus titan, Dasyurus (= Sarcophilus) laniarius, and
Phascolomis mitchelli. It is interesting to note that the common wombat still
existing in eastern Australia was first recognized and named from a fossil skull
and jaw found in the Wellington Caves.

Owen’s interest in the fossil mammals of Australia continued throughout his
long life, and we owe to him a magnificent series of papers published in the
Philosophical Transactions of the Royal Society of London and elsewhere, which
were afterwards collected in his classic Hztinct Mammals of Australia, published
in 1877 (Owen, 1877). There are few cases wherein the work of one man has
contributed so largely to the elucidation of the extinct fauna of any country as
is exemplified in Owen’s researches on the fossil mammals of Australia, and
every subsequent worker has found these volumes a mine of information. The
work is superbly illustrated, most of the specimens being represented of natural
size. The style is dignified, if somewhat ponderous, as was fashionable in the
spacious days of Victoria’s reign; thus Owen calls the rat a “nocturnal murine
omnivorous rodent’.

Australian vertebrate palaeontology undoubtedly owes a great deal to the
enthusiasm, energy and genius of Owen, who was always exhorting his various
friends and correspondents in Australia to procure for him more and better

PRESIDENTIAL ADDRESS. xiii

specimens, so that he succeeded in accumulating in the British Museum, of
which he was Director from 1856 to 1883, a fine series of Australian fossil
vertebrates,

In 1867 Owen wrote to the Colonial Secretary of New South Wales (after-
wards Sir John Robertson) urging that the Government should place a sum of
£200 or £300 on the Estimates for the systematic exploration of the Wellington
Caves. In 1869 an amount of £200 was voted by Parliament for this purpose, and
the work was delegated to the Trustees of the Australian Museum and carried
out under the direction of Gerard Krefft, Curator, and Dr. A. M. Thomson, of
the University of Sydney. As a result over 1,000 specimens were obtained and
forwarded to Owen. Later, in 1881, under the leadership of the late Dr. HE. P.
Ramsay, who succeeded Krefft as Curator of the Australian Museum, additional
specimens were obtained from the caves. A large proportion of the specimens
now forming the collection of Australian fossil mammals in the British Museum
was obtained as the result of these explorations (Caves, 1870, 1882), but a
considerable number were also presented by Dr. George Bennett, his son, G. F.
Bennett, Sir Daniel Cooper, and others.

Some Giants of the Past.

The largest known marsupial, living or extinct, is Diprotodon, a genus
established by Owen on a specimen from the Wellington Caves, consisting of
the anterior portion of the right ramus of the lower jaw containing an incisor
tooth (Owen, in Mitchell, 1838, p. 362, Pl. 31, figs. 1-2); this is the specimen
which had been conjectured to belong to the dugong. Later, in 1843, Owen had
submitted to him a femur and part of a lower jaw with molar teeth, obtained
by Mitchell from the Darling Downs, Queensland. Misled by the character of
the molars, Owen supposed these to belong to a proboscidian to which he gave
the name Dinotherium australe (Owen, 1843a, 18430). But in the following year
he obtained better specimens of the lower jaw, with both incisor and molar teeth,
and recognized that these specimens all belonged to Diprotodon (Owen, 1844).
This was not the only occasion on which Owen was led to predicate the former
existence of proboscidians in Australia, for in the last communication referred
to he described, under the provisional name Mastodon australis, a molar tooth
submitted to him by Count Strzelecki (Owen, 1844, p. 271). This tooth was said
to have been purchased by Strzelecki from a native at Boree Station, Liverpool
Plains, New South Wales, the native stating that further in the interior were
to be found similar and even larger specimens, but that it was impossible for
him to return there owing to the hostility of the tribe upon whose ground the
bones were found (Strzelecki, 1845, p. 312). This was afterwards shown by
Falconer (Falconer, 1868, ii, pp. 271-6) to be the tooth of a South American
Mastodon, M. cordillerum Cuvier (= Cordillerion andium (Osborn) ), and there
can be no doubt that it had been obtained by Strzelecki in his earlier travels in
South America and was not actually the tooth purchased from the native at
Boree. Again, in 1882, Owen described from the Darling Downs a fragment of
a proboscidian tusk under the name Notelephas australis (Owen, 1882). In this
case, too, one surmises that some mistake has been made, and Mr. H. A. Longman,
Director of the Queensland Museum, than whom no one is better acquainted with
the fossils of the Darling Downs, is of opinion that the tusk was not collected
in Australia (Longman, 1916, p. 83).

xiv PRESIDENTIAL ADDRESS.

For thirty years Owen patiently pieced together the available data regarding
Diprotodon before he attempted a reconstruction of the animal, and then it was
incomplete, for the foot structure was still unknown to him. He writes: “Of no
extinct animal of which a passing glimpse, as it were, had thus been caught,
did I ever feel more eager to acquire fuller knowledge than of this huge Marsupial.
No chase can equal the excitement of that on which, bit by bit, and year after
year, one captures the elements of reconstructing the entire creature of which
a single tooth or fragment of bone may have initiated the quest” (Owen, 1877,
p. 190).

Owen died on 18th December, 1892, and earlier in that same year a new
and extensive deposit of Diprotodon bones was discovered in the bed of Lake
Callabonna, South Australia (Stirling, 1894). Here more or less complete skeletons
of Diprotodon and other animals were found and from the fine descriptions by the
late Sir E. C. Stirling and the late A, H. C. Zietz we now know the skeleton of
Diprotodon in almost every detail (Stirling and Zietz, 1899), and most of the
large museums of the world now possess casts of this interesting animal,
constructed at the South Australian Museum. These show that Diprotodon was
as large as the largest rhinoceros.

While we are on Diprotodon, I might mention an interesting piece of history
regarding one of the most important and best known finds of its remains, namely,
the fine skull now in the British Museum (Owen, 1877, Pl. 19, figs. 1-4), of which
the story seems to be as follows: This skull was found at King’s Creek, Darling
Downs, Queensland, and was, with other specimens, brought to Sydney in 1847
by a Mr. Turner. Here it was put together by Ludwig Leichhardt, the celebrated
explorer, who, as is well known, perished in an attempt to cross Australia from
east to west, the Rev. W. B. Clarke, the “Father of Australian Geology”, and
W.S. Wall. Curator of the Australian Museum. This collection was subsequently
acquired by Benjamin Boyd, founder of Boydtown and of the whaling industry
of Twofold Bay, New South Wales, who afterwards disappeared into the Pacific
and is believed to have been killed by the natives of Guadalcanar. The collection
was forwarded by Boyd to England, but the vessel carrying it was wrecked off
Beachy Head. The skull was, however, recovered and was sold with other
specimens to the British Museum (Leichhardt, 1867, p. 63, footnote; Clarke, 1853a,
p. 5). One interesting feature of this skull was referred to by W. B. Clarke,
namely, that it “was in so apparently fresh a condition when it was first disen-
tombed, that the brain was in a state to be easily recognized, and traces of
blood vessels in it were distinct” (Clarke, 1853b, p. 12). This would suggest
that Diprotodon was alive at no distant date, geologically speaking.

It is interesting to speculate on the possibility of Diprotodon having survived
until the coming of man to Australia, and even of some traditions among the
aborigines regarding the former existence of these animals. I have come across
two references to this possibility and there may be others unknown to me.
One occurs in an interesting paper by Leichhardt, entitled “Notes on the Geology
of Parts of New South Wales and Queensland made in 1842-438”, which was
published in Germany, translated by G. H. F. Ulrich and edited by W. B. Clarke
for the Australian Almanac, 1866-67 (Leichhardt, 1855, 1867). In it occurs the
following passage, referring to bones of a gigantic animal found in the Darling
Downs; these Leichhardt describes as similar to bones from the Wellington
Valley which were at first taken for elephant bones, but which he was firmly
convinced were wrongly determined, holding that these animals, “like all the

PRESIDENTIAL ADDRESS. XV

accompanying smaller animals, were formed after the Australian type, and
belong, probably, exclusively to this continent”. Leichhardt goes on to say:
“It is possible that it is at the present time still living in the tropics of this
country, which are rich in water. This I was told by Mr. Dennis, that the
blacks speak of extensive lakes and gigantic animals” (1855, pp. 38-39; 1867,
pp. 62-63).

Dr. George Bennett is more explicit (Bennett, 1872, p. 315), writing: “I have
had a long conversation with ‘Charlie Pierce’, an aboriginal, relative to these
fossils; and he avers that they are those of an animal, long extinct, known to
the natives by the name ‘gyedara’. Tradition among them has handed down
the appearance and habits of the animal for generations, but Charlie says he
never paid much attention to the descriptions that have been given to him, but
imagines the animal was as large as a heavy draught horse, walked on all fours
the same as any other four-footed beast, eating grass, never went any distance
back from the creeks to feed, and spent most of its time in the water, chiefly
in enormous holes excavated in the banks. I told him he must mean some
other animal; but he spoke most positively and asserted that the bones which we
have been finding are those of the animal of which he was speaking, and that
at one time the bones were very numerous about the Gowrie waterholes, where
his forefathers had seen the animals themselves sporting about. I again asked
him if they did not live on the leaves of trees, and his reply was that they were
never seen to feed on them but always on grass, the same as a horse or a
bullock.”

We should, of course, make allowance for the difficulty of getting reliable
information from the aborigines, and their proneness to give the replies which
they expect are desired, but Dr. Bennett, who was an excellent observer and
investigator, seems to have exercised due discretion in making his enquiries,
and it seems possible that the aborigines did have legends regarding the former
existence of some large mammal.

Another interesting large marsupial genus of the Australian Pleistocene is
Nototherium, which resembles Diprotodon in some respects and is usually included
in the same family. It was first made known by Owen in 1845 (Owen, 1845,
p. 314), and a fine skull, now in the Australian Museum, was discovered in 1856,
at King’s Creek, Darling Downs; this is the specimen which was given the name
Zygomaturus trilobus by William Sharp McLeay, but this is a synonym of
Nototherium mitchelli Owen. In more recent years practically complete skeletons
have been found in lake and swamp areas in Tasmania and King Island and,
thanks to the excellent descriptions by Messrs. H. H. Scott and C. E. Lord (Scott,
1915; Scott and Lord, Pap. Proc. Roy. Soc. Tas., 1920-1924), the structure and
appearance of the animal are now well known. It was about as large as a bullock
and was remarkable for the depth and the width of its skull.

In the nototherian genus Huryzygoma of Longman (1921) the width of the
skull is greater than its length.

Another interesting’ extinct form from the Australian Post-Tertiary is
Thylacoleo carnifex Owen, the so-called Marsupial Lion, which is distinguished
above all by its huge sectorial premolars. These teeth and also the incisors
were figured in Mitchell’s Three Hapeditions (Mitchell, 1838, Pl. 32, figs. 4-11),
but no mention of them is made in the text. The name Thylacoleo first appeared
in Gervais’ Zoologie et Paléontologie francaises, 1st edition, 1848-1852, p. 192,
and the species was described by Owen in 1859 (Owen, 1859, p. 309; 1877, p. 107).

xvi PRESIDENTIAL ADDRESS.

Only the skull and mandible of this singular animal are known with certainty,
although certain limb bones have been tentatively assigned to it. The skull is
rounded, about as large as that of a leopard, and the presence of pointed incisors
and large sectorial premolars, the reduction of the molars, and the general
structure of the skull and mandible, led Owen to believe it to be a carnivore,
indeed “one of the fellest and most destructive of predatory beasts’ (Owen,
1877, p. 119). That it was carnivorous in habit, which was disputed by Flower
(1868) and Krefft (1866a; 1872; 1873, p. 188) but strongly supported by Broom
(1898), is now the generally accepted opinion, and I myself seem to be the only
one who still has doubts (Anderson, 1929, pp. 44-47).

Brief mention may be made here of the curious ungual phalanges found in
the Wellington Caves and elsewhere, which led Krefft to surmise that an animal
related to the sloths of South America, and to which he gave the name Mylodon (7?)
australis (Krefft, in Caves, 1870, p. 7), once inhabited Australia. Etheridge has
discussed these phalanges exhaustively (Etheridge, 1918), and concludes that there
were two kinds, one of which perhaps belonged to Thylacoleo, the other possibly
to a large extinct phalanger.

These large extinct marsupials of the Australian Post-Tertiary illustrate
what has been frequently observed, namely, that just before extinction there is
a tendency to an increase of bulk over what is usual for the group to which
an animal belongs. Large size is a kind of specialization which seems to
handicap the race, especially if from some change in the environment, food
supplies become limited. It is generally supposed that when Diprotodon and
its contemporaries roamed over Australia our climate was more humid and
that extensive freshwater lakes and abundant vegetation existed where now we
have, in part, desert country. But Darwin has maintained (1884, pp. 85-89) that
the assumption that large animals require a luxuriant vegetation is completely
false and has vitiated the reasoning of geologists on some points of great
interest in the ancient history of the world. He instances South Africa, which
is a sterile country, yet it supports an extraordinary number of animals, some
of large bulk. It does not necessarily follow, therefore, that during what we
may call the Diprotodon period, the climate and vegetation of Australia were
very different from what they are to-day.

The Macropodidae.

The majority of the known fossil mammals of Australia belong to this family,
still the most largely represented, and comprising the kangaroos, wallabies, and
rat-kangaroos. Most of the extinct forms, which have been described chiefly by
Owen (1877), or by De Vis (1894), are very similar to living species.

The largest was Palorchestes, of which the skull, about sixteen inches in
length, is comparable with that of an ox. Only portions of the skull and
mandible, and girdle and limb bones of this large kangaroo are known. It was
by far the largest of the kangaroos, but from the structure of its limb bones
it is conjectured that it progressed by leaps as its living relatives do. Its
lower molars, while of the macropodine type, present some resemblance to those
of Nototherium and Diprotodon, but its lower incisors were procumbent and
spatulate, as in typical kangaroos.

The two extinct genera, Procoptodon and Sthenurus, which De Vis (1894,
p. 88) has united under the name Sthenuwrus, are closely related to one another
and distinguished from Macropus by skull and tooth structure. The skull was
relatively short and deep, in general shape recalling that of the koala

PRESIDENTIAL ADDRESS. xvii

(Phascolarctus) (Anderson, 1932). In some species of Procoptodon in particular,
the molar teeth are complex in structure with numerous folds and buttresses,
and the lower incisors are sub-cylindrical and semi-erect, not spatulate and
procumbent as in existing kangaroos; in these respects the lower incisors of
Procoptodon suggest comparison with those of Nototheriwm.

These two extinct forms were evidently powerfully built, stocky animals,
not slender graceful creatures like modern kangaroos and wallabies.

An interesting form, the affinities of which are uncertain, but which may
be an annectant between the Macropodidae and the Phalangeridae is Triclis
oscillans De Vis (1888), renamed Propleopus oscillans by Longman (1924b,
pp. 20-21). The only part known is the lower jaw, which has an additional
tooth, probably a canine, between the premolar and the incisor, such a tooth
not being represented otherwise in the Macropodidae.

Phalangeridae.

Phalangers are not well represented among fossil forms, possibly because
of their generally arboreal habits. The most interesting is Wynyardia bassiana,
from the Table Cape beds of Tasmania, now generally regarded as of Upper
Miocene or Lowest Pliocene age. It is therefore the oldest known Australian
marsupial, and on this account its affinities have been eagerly discussed. The
only known specimen consists of a broken skull and mandible, some damaged
vertebrae and certain bones of the appendicular skeleton; unfortunately no teeth
were found.

This celebrated specimen, now in the Tasmanian Museum, Hobart, was first
referred to by Tenison-Woods in 1877 (1877, pp. 73-74), who describes it as the
almost perfect skeleton of a wallaby, Halmaturus (?). It is also mentioned
by Johnston (1888, pp. 261, 288) as a Halmaturus. In 1900 it was described
and named by Baldwin Spencer (1900), who regarded it as a form in some
respects intermediate between the diprotodont and the polyprotodont marsupials,
allied on the one hand to the Phalangeridae and on the other to the Dasyuridae.
This opinion has not been substantiated by later authors. Thus Bensley (1903,
p. 200) states that the relations of Wynyardia are more probably with the advanced
genera, such as Pseudochirus, Phascolarctus or Phalanger. Gregory (1920, p. 143)
remarks on the lacrymal bone and states that similar types of lacrymal are
to be seen in the modern phalangers and their allies and the Macropodidae.
Osgood (1921, pp. 136-139) classifies it with the diprotodonts as the type of a
distinct family Wynyardiidae, and finds a number of points common to Wynyardia
and Caenolestes. Longman (1924a, pp. 8-9) finds no valid reason for suggesting
that Wynyardia and Caenolestes are related, but does not express an opinion as
to the affinities of the former. Wood (1924, pp. 83-84) regards it as a slightly
primitive phalanger, and Simpson (1930, pp. 7, 65) classes it with the
Phalangeridae. Wood Jones (1930) makes a careful re-examination of the skeleton
and concludes that it is a fully developed and typical Australian diprotodont, in
no way an annectant between the Polyprotodontia and Diprotodontia, that it
shows many resemblances to the living genus TJrichosurus, and must be considered
an ally of the phalangers.

Another interesting form is Burramys parvus Broom (1895, 1896), from a
small bone breccia deposit on the top of a limestone hill near the Wombeyan
Caves, N. S. Wales; the specimens found consisted of some moderately good lower
jaws and fragments of the upper. It is a small animal, the lower jaw less

XVili PRESIDENTIAL ADDRESS.

than half an inch in length, and in its large grooved premolar and other tooth
characters it shows a resemblance to the rat-kangaroos, while the jaw itself is
more phalangerine. Broom (1895, p. 566) suggested that in Burramys we have
one more link in the chain binding the kangaroos and the phalangers, the main
links being Macropus, Aepyprymnus, Hypsiprymnodon, Burramys. In a later paper
(Broom, 1898, p. 63) he says it is evidently the representative of a sub-family
Burramyinae of the Phalangeridae, and that, as a phalanger with posterior
premolars enormously enlarged, it comes nearer to Thylacoleo than does any
extinct or living form hitherto discovered.

From the Pleistocene of Queensland De Vis (1889) has described fragmentary
remains as those of large members of this family, to which he gives the names
Koalemus ingens (pp. 106-9), Archizonurus securus (pp. 109-111), and Cuscus.
procuscus (pp. 111-112).

Wombats.

A number of phascolomyid species have been found fossil in Australia, and
it has been already mentioned that Phascolomis mitchelli was first described
from a Wellington Caves specimen. The largest known member of the family is
Phascolomis gigas Owen, which was widely distributed. Specimens found at
Lake Callabonna enabled Stirling (1913, pp. 127-178) to describe a large part
of its skeleton; he estimates that it was about twice as large as any existing
wombat and had numerous points of resemblance to Diprotodon (loc. cit., p. 177).

Polyprotodont Marsupials.

The list of polyprotodont marsupials found fossil in Australia is not a
long one, and no new genera or outstanding forms have been discovered, the
species being either identical with those still living or very closely related to these.

Fragments of jaws and teeth of the family Peramelidae have been found in
the Wellington Caves and elsewhere (Owen, 1877, p. 107; Broom, 1896, p. 56;
Glauert, 1926, p. 70), but it is doubtful whether any of the fossil forms are
distinct from living species, and no special interest attaches to any of the finds.

The most important feature regarding the fossil dasyurids is that the two
genera Thylacinus and Sarcophilus, now restricted to Tasmania, formerly existed
on the mainland as well. It is generally supposed that on the mainland they
were exterminated by the dingo, which never got into Tasmania. The mainland
forms were very similar to the existing Tasmanian species, but Thylacinus
spelaeus Owen and Sarcophilus laniarius Owen seem to have been rather larger
than the island forms. An almost complete skeleton of a thylacine, scarcely, if
at all, separable from the living Thylacinus cynocephalus, has been found in the
Jenolan Caves, and is now in the collection of the Department of Mines, N. 8S.
Wales. The bones are brittle, but their condition indicates no great age, and it
would appear that the extinction of Thylacinus on the Australian Continent
took place comparatively recently.

Monotremes.

The geological history of this, the most interesting mammalian Order, is prac-
tically unknown, and no signs of monotremes have been found elsewhere than in
Australia, although more than once the discovery of supposed members of the
Order in other countries has been announced. The alleged occurrence of mono-
tremes in the Santa Cruz beds of South America (Ameghino, 1894, pp. 182-9)
is based on an erroneous determination, the bones being really those of aberrant

PRESIDENTIAL ADDRESS. xix

armadillos (Lydekker, 1894, p. 67). So, too, Xenotherium wunicum, from the
Oligocene of Montana, described by Douglass (1905) as a possible monotreme,
has been shown by Matthew (1928), Zdansky (1927) and Simpson (1927) to
be an edentate. Abel has recently advanced the opinion that Desmostylus,
usually classed in the Sirenia, is a descendant of the Multituberculata and
ancestral to the monotremes (in Weber, 1928, pp. 48-44), but this opinion has
not been generally accepted (Simpson, 1928b, p. 182; 1929, p. 12). In fact,
nothing is known of the history of the monotremes prior to the Pleistocene or late
Pliocene of Australia, and the specimens found there are very similar to existing
species and do not enable us to do more than guess regarding the origin and
evolutionary history of these mysterious animals. It has more than once been
suggested that the monotremes are descended from or related to the Multi-
tuberculata (Cope, 1888; Broom, 1914), largely on account of the supposed
similarity of the teeth of Ornithorhynchus to the molars of the multituberculates.
Simpson (1929) has recently made a careful study of the dentition of
Ornithorhynchus, and considers that the evidence of the teeth is strongly against
the view that the platypus is related to the multituberculates, and emphasizes
that the teeth of Ornithorhynchus are not like those of any other known animal
in form, and that, though a vague resemblance to those of the Triconodonta
may eventually prove to be significant, this at present is not trustworthy. Nor
does he find any similar dentition among mammal-like reptiles.

It seems possible that the monotremes may never have existed outside
Australia, and that they have evolved from the Reptilia independently of the
rest of the Mammalia, although Gregory (1910, p. 160) has suggested the
possibility that the common ancestors of monotremes, marsupials and placentals
were already mammals.

The occurrence of fossil monotreme remains in Australia has been summarized
by Dun (1895). Krefft (1868) was the first to announce the discovery of a
monotreme fossil, when he described the articular head of a humerus from the
Pleistocene of Queensland under the name *Hchidna oweni. Owen (1884) described
from the Wellington Caves the left humerus of an echidna, probably identical with
Krefft’s species. In 1870 Krefft mentioned the occurrence in the Wellington
Caves of a “fractured femur of a species larger than our present Hchidna hystriz”’
(Krefft, in Caves, 1870, p. 10). In 1885 De Vis (1885) announced the first dis-
covery of fossil remains of Ornithorhynchus, describing a mandible and tibia of a
small species, Ornithorhynchus agilis, from the Pleistocene of the Darling Downs.
Dun (loc. cit.) added two new species, Ornithorhynchus maximus and Echidna
robusta, from the Canadian Lead, Gulgong, N. S. Wales, generally regarded as
of Pliocene age. Of the former a right humerus was found, of the latter portion
of the skull and an atlas vertebra. Later Broom (1896, p. 58) discovered near
the Wombeyan Caves a number of bones of a large echidna, comprising portion of
the ilium and sacrum, the distal end of a left femur, the articular head of a
left femur, two vertebral centra, and a number of fragmentary long bones; these
he considered to be ascribable to Echidna oweni Krefft. Glauert described the
right humerus of Tachyglossus aculeatus (1910, p. 18-14), and the atlas vertebra,
the clavicles and episternum, the pelvic girdle, two femora, a tibia, and a
radius of a new species, Zaglossus hacketti, twice as large as the living Tachy-
glossus aculeatus (Glauert, ibid., pp. 244-8). Scott and Lord (1921a) have recorded
the right femur and portion of the proximal end of a humerus of a new species,

* The generic name Hchidna is preoccupied and should be replaced by Tachyglossus.

xX PRESIDENTIAL ADDRESS.

Zaglossus harrisoni, from King Island, Tasmania; this species was fully twice as
large as Tachyglossus aculeatus. Weber (1928, p. 43) refers to Echidna amplor
De Vis, but I have not been able to trace the original description.

This completes the record of described monotreme fossil remains, and reference
to the descriptions shows that, while several of the extinct species are consider-
ably larger than their living relatives, there are few and unimportant structural
differences.

Rodents.

Of placental mammals the most abundant of the fossil forms found in
Australia are rodents, of which numerous jaws and other fragments have been
found in the Wellington Caves, but these have not been systematically examined,
although Krefft states that this deposit contains six or more species of Hapalotis
and Mus which may be distinguished from the living forms (Krefft, 1871, p. 715).
Broom (1896, p. 59) also refers to innumerable remains of Bush Rats (Mus sp.)
in a bone breccia near the Wombeyan Caves.

Cetacea.

Fossil cetaceans are known from the Tertiary of southern Australia, the
specimens found consisting mainly of teeth. McCoy (1867) described a tooth
from Cape Otway, Victoria, as Squalodon wilkinsoni, a name later changed to
Parasqualodon Hall (1911, p. 262). A molar tooth from Wellington, South
Australia, was named Zeuglodon harwoodi by Sanger (1881) (= Metasqualodon,
Hall, loc. cit., p. 263). The skull and parts of the skeleton of another squalodont
whale, from Table Cape, Tasmania, were recorded by Flynn (1920, 1923) as
Prosqualodon davidis; it belongs to a short-beaked group of the Squalodontidae
which is also represented in Patagonia.

Chapman has described a delphinid Steno cudmorei from the Kalimnan of
Beaumaris, Victoria (1918a, pp. 41-42), and Scott and Lord (19210) have recorded,
from Wynyard, Tasmania, delphinoid vertebrae and portions of a skull, which,
they suggest, may belong to the genus Globicephalus.

Of the physeteroid whales several species have been found fossil. McCoy
(1879, pp. 19-20) described a new genus and species, Physetodon baileyi, from
Mordialloc, Victoria, and Chapman has described two species of Scaldicetus,
namely S. macgeei from Beaumaris (1912) and S. lodgei from the Oligocene of
Muddy Creek, Victoria (1918a, pp. 34-35). A new genus and species of physeteroid
whale, Scaptodon lodderi, was founded by Chapman (19180) on a tooth washed
up twenty-eight miles east of Table Cape, Tasmania, and Scott (1913) has
described the arm bones and vertebrae of a supposed ziphioid whale from Wynyard,
Tasmania.

The Dingo.

Much has been written regarding the dingo and its coming to Australia, some
authorities claiming it as a member of the indigenous fauna of Australia, others
believing that it was introduced by the aborigines. According to Wood Jones
(1920), the dingo is a true dog, related to the northern wolf, and it cannot,
therefore, be indigenous to Australia. Yet there is evidence that it has long
been an inhabitant, and its teeth have been found in association with the
remains of extinct marsupials (Etheridge, 1916a), and it is believed, as stated
previously, that the dingo was responsible for the extermination on the mainland
of Thylacinus and Sarcophilus. The difficulties of the hypothesis that the dingo

PRESIDENTIAL ADDRESS. xxi

came overland are very great, and Wood Jones (loc. cit.) confidently asserts
that the dingo came by boat along with the aborigines.

Man.

If man brought the dingo into Australia and the dingo was contemporaneous
with some of the extinct marsupials, then we must suppose that man has a fairly
long geological history in Australia, a subject which has also been largely
discussed. Etheridge (1890, 19160) has carefully reviewed the evidence, and
David (1923) has more recently contributed an important paper on this subject.
One of the best pieces of evidence as to the early existence of man in Australia
is the portion of a human tooth found by Krefft in the bone breccia of the
Wellington Caves. Krefft (1874) definitely states that this was found in the
solid breccia, and there is no reason to doubt his word. I sought confirmation of
this from the late Dr. J. Mildred Creed, who informed me that he was present
when the tooth was found. Krefft also mentions (in MS.) finding parts of a
human skeleton in the Wellington Caves (Etheridge, 19160), which may be the
bones described in his “Australian Vertebrata, Recent and Fossil” (Krefft, 1867,
p. 49). It is, however, very probably the case that not all bones found in the
Wellington Caves are of the same age, though there is strong evidence that man
has a considerable antiquity in Australia.

Conclusion.

It will, I think, be realized that in little over a hundred years much has
been accomplished in the investigation of the fossil mammals of Australia, yet
there are still immense gaps in our knowledge. Information regarding the
Tertiary marsupials is almost a blank, the only form definitely known to be of
that age being Wynyardia, a fully developed phalanger. It is true that some of
the supposed Pleistocene mammals may really be Tertiary and others Holocene.
It is certain, however, that there must have been marsupials in Australia through-
out the Tertiary period and that their deployment was accomplished in Australia
itself, and such forms as Diprotodon, Nototherium and Thylacoleo must have had
a long line of ancestors. Yet, so far as actual evidence is concerned, they seem
to have sprung into existence fully developed, in fact, over developed and ripe
for extinction. The same is the case with the monotremes, which probably have
a longer history in Australia than the marsupials. It is not beyond hope,
however, that Tertiary mammals may yet be found here in some abundance, and
that the evolutionary lines and inter-relationships of our mammals may be
discovered, but it seems that organized and diligent search and considerable
expenditure will be necessary. It is likely that the earliest Australian marsupials
were small mouse-like creatures the bones of which might easily escape preserva-
tion and might easily be overlooked during mining or well-sinking operations.

The place of origin and the time and mode of entry into Australia of our
marsupials is still unknown. Dr. G. G. Simpson (1930, pp. 6-7), one of the latest
writers to discuss this question, considers that much evidence points to the
Cretaceous as the time of entry, and that as regards the immigration route, a
connection of Australia and South America with Holarctica, but not directly
with one another, affords the simplest and least hypothetical solution. Unfor-
tunately no marsupials have yet been found in Asia, although primitive insectivora
have been described from the Cretaceous of Mongolia (Gregory and Simpson,
1926; Simpson, 1928a, 1928c).

xxii PRESIDENTIAL ADDRESS.

It is interesting to note that a supposed marsupial, Palaeothentoides africanus,
which shows affinities with the Caenolestidae of South America, has recently been
described from the Middle Pliocene of Little Namaqualand, South Africa (Stromer,
1931). Dr. Stromer, in discussing its possible origin, suggests that there may have
been some early migration route from South America to Africa, or that this
marsupial may be independently derived from some early Huropean marsupials.
Dr. G. G. Simpson, however (1933), is inclined to doubt whether this really is
a marsupial and, even if it is, whether it has any relationship to the South
American caenolestids.

Apart from the lack of Tertiary marsupials in Australia, it cannot be said
that our Pleistocene forms are yet well known. Many of the specimens found
are fragmentary, consisting of portions of skulls and jaws, and unassociated
bones of the skeleton, complete skeletons being rarely found. It is hoped, however,
that as time goes on better specimens with all the bones associated may be
found so that we shall have a better idea of the appearance in life of Thylacoleo,
Palorchestes, Procoptodon and others.

List of Works Referred to.

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Bocéne de Patagonie. Bol. Acad. Nac. Cien. Cordoba, i, 13, pp. 259-452; also
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ANDERSON, C., 1929.—Palaeontological Notes, No. I. Macropus titan Owen and Thylacoleo
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, 1932.—Palaeontological Notes, No. III. The skull of Sthenurus atlas Owen.
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BENNETT, G., 1872.—A Trip to Queensland in Search of Fossils. Ann. Mag. Nat. Hist.,
(4), Ix, pp. 314-321.

BENSLEY, B. A., 1903.—On the Evolution of the Australian Marsupialia; with remarks
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Broom, R., 1895.—On a small fossil Marsupial with large grooved premolars. Proc.
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, 1896.—Report on a bone breccia deposit near the Wombeyan Caves, N.S.W.;
with descriptions of some new species of Marsupials. Jbid., xxi, pp. 48-61.

, 1898.—On the affinities and habits of Thylacoleo. Ibid., xxiii, pp. 57-74.

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Caves, Wellington (Correspondence relative to Exploration of), 1870.—Parliamentary
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Caves and Rivers of New South Wales, Exploration of, 1882.—Parliamentary Papers.
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CHAPMAN, F., 1912.—On the occurrence of Scaldicetus in Victoria. Rec. Geol. Surv.
Vict., iii, pp. 236-8.

, 1914.—Australasian Fossils. George Robertson & Co., Melbourne.

,1918a.—Some Tertiary Cetacean remains. Proc. Roy. Soc. Vict. (N.S.), Xxx,
pp. 32-438.

,1918b.—On an apparently new type of Cetacean tooth from the Tertiary of
Tasmania. Ibid., pp. 149-152.

CLARKE, W. B., 1853a.—(On the Geology of the basin of the Condamine River.) Rept. X,
14th Oct., Papers relative to Geol. Surveys, N. S. Wales, pp. 1-11.

, 1853b.—Extract from remarks on the bones brought to Sydney by Mr. Turner.
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Cope, E. D., 1888.—The Multituberculata Monotremes. Amer. Nat., xxii, p. 259.

Darwin, C., 1884.—Journal of Researches. London, John Murray.

, 1886.—The Origin of Species. London, John Murray.

Davip, T. W. E., 1923.—Geological Evidence of the Antiquity of Man in Australia
(Johnston Memorial Lecture). Pap. Proc. Roy. Soc. Tas., pp. 109-150.

PRESIDENTIAL ADDRESS. xxiii

Dr Vis, C., 1885.—On an Extinct Monotreme, Ornithorynchus agilis. Proc. Roy. Soc.
Q’land, ii, pp. 35-38.

,1888.—On an Extinct Genus of the Marsupials allied to Hypsiprymnodon.
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, 1889.—On the Phalangeridae of the Post-Tertiary period in Australia. Proc.
Roy. Soc. Q’land, vi, pp. 105-114.

, 1894.—A review of the fossil jaws of the Macropodidae in the Queensland
Museum. Proc. Linn. Soc. N.S.W., (2), x, pp. 75-133.

Douaeuass, E., 1905.—The Tertiary of Montana. Mem. Carnegie Mus., ii, 1905-1906,
pp. 204-8.

Dun, W. S., 1895.—Notes on the occurrence of Monotreme remains in the Pliocene of
New South Wales. Rec. Geol. Surv. N. S. Wales, iv, pp. 118-126.

ETHERIDGE, R., JuN., 1878.—A Catalogue of Australian Fossils. Cambridge Univ. Press.

,1890.—Has man a geological history in Australia? Proc. LINN. Soc. N.S.W.,
(2), v, pp. 259-266.

, 1916a.—The Warrigal or “Dingo”, introduced or indigenous? Mem. Geol. Surv.
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, 1916b.—Occasional Notes. No. I. Antiquity of Man in Australia. Rec. Austr.
Mus., xi, pp. 31-32.

,1918.—The Ungual phalanges terraed Mylodon australis by Krefft, spelaean
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(9), ii, pp. 307-318.

FALCONER, H., 1868.—Palaeontological Memoirs, ii, pp. 271-6. London.

FLOWER, W., 1868.—On the affinities and probable habits of the extinct Australian
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, 1923.—A whale of bygone days. Austr. Mus. Mag., i, pp. 266-272.

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, 1920.—Studies in Comparative Anatomy and Osteology; No. IV. A review of
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,and SIMPSON, G. G., 1926.—Cretaceous mammal skulls from Mongolia. Amer.
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Hau, T. S., 1911.—On the systematic position of the species of Squalodon and
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, 1871.—Australian Vertebrata (fossil and recent). Industrial Prog. N. 8S. Wales,
Intercol. Exhib., 1870, at Sydney, pp. 699-780; also separate, pp. 96.

, 1872.—A Cuvierian principle in palaeontology, tesi'ed by evidences of an extinct
leonine marsupial (Thylacoleo carnifex). Ann. Mag. Nat. Hist., (4), x, pp. 169-182.

, 1873.—Mammals of Australia and their classification. Trans. Roy. Soc. N. S.
Wales, pp. 135-147.

KKREFFT, G., 1874.—Further discovery of the remains of a great extinct wingless bird in
Australia. Geol. Mag., i, pp. 46-47.

XXiV PRESIDENTIAL ADDRESS.

Lane, J. D., 1831.—Account of the discovery of bone caves in Wellington Valley, about
210 miles west from Sydney in New Holland. Hdinb. New Phil. Journ., pp. 364-368.

LEICHHARDT, L., 1855.—Beitrage zur Geologie von Australien. Abh. nat. Ges. Halle,
iii, pp. 1-62.

, 1867.—Notes on the geology of parts of New South Wales and Queensland.
Australian Almanac, pp. 59-80 (Transl. of 1855).

LONGMAN, H. A., 1916.—The supposed artiodactyle Queensland fossils. Proc. Roy. Sec.
Q’land, xxviii, p. 83.

, 1921.—A new genus of fossil marsupials. Mem. Q’land Mus., vii, pp. 65-80.

, 1924a.—The Zoogeography of Marsupials. TJIbid., viii, pp. 1-15.

, 1924b.—Some Queensland fossil vertebrates. JIbid., pp. 16-28.

LYDEKKER, R., 1894.—Contributions to a knowledge of the fossil vertebrates of
Argentina. Ann. Mus. La Plata, iii, pp. 66-70.
MatTrHEW, W. D., 1928.—Xenotherium an edentate. Journ. Mamm., ix, pp. 70-71.
McCoy, F., 1867.—On the occurrence of the Genus Squalodon in the Tertiary strata of
Victoria. Geol. Mag., iv, p. 1465.
, 1879.—Prodromus of the Palaeontology of Victoria. Decade vi, pp. 19-20.
MITCHELL, T. L., 1831.—An account of the limestone caves at Wellington Valley, and of
the situation, near one of them, where fossil bones have been found. Proc. Geol.
Soc. Lond., i, pp. 321-2.

, 1838.—Three expeditions into the interior of Eastern Australia, ii, pp. 359-363.
London.

Oscoop, W. H., 1921.—A monographic study of the American marsupial Caenolestes.
Field Mus., Zool. Ser., xiv, pp. 4-162.

OWEN, R., 1843a.—On the discovery of the remains of a mastodontoid pachyderm in
Australia. Ann. Mag. Nat. Hist., xi, pp. 7-12.

, 1843b.—Additional evidence proving the Australian pachyderm .. . to be a
Dinotherium. Ibid., pp. 329-332.

, 1844.—Description of a fossil molar tooth of a mastodon, discovered by Count
Strzelecki in Australia. TIbid., xiv, pp. 268-271.

, 1845.—Descriptive and illustrated catalogue of the fossil organic remains of
Mammalia and Aves contained in the Museum of the Royal College of Surgeons of
England. London.

, 1859.—Description of a mutilated skull of a large marsupial carnivore. Phil.
Trans., cxlix, pp. 309-322.

, 1877.—Researches on the fossil remains of the extinct mammals of Australia,
with a notice of the extinct marsupials of England. Two vols. London.

, 1882.—Description of portions of a tusk of a proboscidian mammal
(Notelephas australis Owen). Phil. Trans., clxxiii, pp. 777-781.

, 1884.—Evidence of a large extinct Monotreme (Hchidna ramsayi Owen) from
the Wellington breccia caves, New South Wales. IJbid., clxxv, pp. 273-275.

PENTLAND, W., 1831.—Further notes in regard to the fossil bones found in Wellington
Country, New South Wales. Edinb. New Phil. Journ., pp. 179-180.

, 1832.—On the fossil bones of Wellington Valley, New Holland or New South
Wales. Ibid., pp. 301-308.

, 1833.—Observations on a collection of fossil bones sent to Baron Cuvier from
New Holland. Ibid., pp. 120-121.

Sancer, E. B., 1881.—On a molar tooth of Zeuglodon from the Tertiary beds on the
Murray River, near Wellington, S.A. Proc. LInn. Soc. N.S.W., v, pp. 298-300.
Scott, H. H., 1913.—Notes on a fossil whale from Wynyard, Tasmania. Pap. Proc. Roy.

Soc. Tas., pp. 167-172.

, 1915.—A monograph of Nototherium tasmanicum. Rec. Geol. Surv. Tas., No. 4.

and Lord, C. E., 192la.—Studies in Tasmanian mammals, living and extinet.
PGpe PrOCch ROY ISOCy Lasy, PDs) Leo.

, 1921b.—Studies in Tasmanian mammals, living and extinct. Jbid., pp. 180-181.

Simpson, G. G., 1927.—A North American Oligocene edentate. Ann. Carnegie Mus., xvii,

pp. 283-296.
,1928a.—Further notes on Mongolian Cretaceous Mammals. Amer. Mus.
Nov., 329.

, 1928b.—A catalogue of the Mesozoic Mammalia in the Geological Department
of the British Museum, p. 182. London. British Museum.
Simpson, G. G., 1928c.—Affinities of the Mongolian Cretaceous Insectivores. Amer. Mus.
Nov., 330.

PRESIDENTIAL ADDRESS. XXV

Simpson, G. G., 1929.—The dentition of Ornithorhynchus as evidence of its affinities.
Tbid., 390.
, 1930.—Post-Mesozoic Marsupialia. Fossilium Catalogus, Animalia, pars 47.
Berlin, W. Junk. ,
— ,1933.—The supposed fossil marsupial from Africa. Nat. Hist., xxxiii, pp. 106-7.
Spencer, B., 1900.—A description of Wynyardia bassiana, a fossil Marsupial from the
Tertiary beds of Table Cape, Tasmania. Proc. Zool. Soc. Lond., pp. 776-795.
STIRLING, FE. C., 1894.—The recent discovery of fossil remains at Lake Callabonna,
South Australia. Nature, 1, p. 186.

—,1913.—Fossil remains of Lake Callabonna, Part iv, 2. On the identity of
Phascolomys (Phascolonus) gigas Owen and Sceparnodon ramsayi Owen, with a
description of some of its remains. Mem. Roy. Soc. S. Austr., i, pt. iv, pp. 127-178.

and Zinrz, A. H. C., 1899.—Fossil remains of Lake Callabonna. Pt. i. Descrip-

tion of the manus and pes of Diprotodon australis. Ibid., i, Pt. i, pp. 1-40.

StrRoMER, E., 1931.—Palaeothentoides africanus, nov. gen., nov. spec., Hin Erstes
Beuteltier Aus Afrika. Site. Bayer. Akad. Wiss. Math.-Naturw., Abt. 3, pp. 177-190.

SvRZELECKI, P. BE. ps, 1845.—Physical description of New South Wales and Van
Diemen’s Land. London.

WEBER, M., 1928.—Die Si&ugetiere, II, 2nd Edition. Jena.

Woop, H. E., 1924.—The position of the “Sparassodonts’”, with notes on the relation-
ships and history of the Marsupialia. Bull. Amer. Mus. Nat. Hist., li, pp. 77-101.

Woops, J. E. T., 1877.—On the Tertiary deposits of Australia. Journ. Proc. Roy. Soc.
N. S. Wales, xi, pp. 65-82.

ZDANSKY, O., 1927.—Ueber die Systematische Stellung von Xenotherium Douglass. Bull.
Geol. Inst. Univ. Upsala, xx, pp. 231-6.

Dr. G. A. Waterhouse, Honorary Treasurer, presented the balance sheets for
the fourteen months ended 28th February, 1933, duly signed by the Auditor,
Mr. F. H. Rayment, F.C.A. (Aust.); and he moved that they be received and
adopted, which was carried unanimously.

No nominations of other Candidates having been received, the President
declared the following elections for the ensuing Session to be duly made:

President: Professor A. N. St. G. H. Burkitt, M.B., B.Sc.

Members of Council: C. Anderson, M.A., D.Sc., Prof. A. N. St. G. H. Burkitt,
Meee bs SC.n Hey Jn Carter, BEAL WES: Prot. We J. Dakins DSc, A. G. Hamilton,
Prof. J. Macdonald Holmes, B.Sc., F.R.G.S., A. F. Basset Hull, and C. A.
Sussmilch, F.G.S.

Auditor: F. H. Rayment, F.C.A. (Aust.).

A cordial vote of thanks to the retiring President was carried by acclamation.

XXVi

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THE MAYFLIES OF THE MOUNT KOSCIUSKO REGION. I.
(PLECTOPTERA.)

INTRODUCTION AND F'AMILY SIPHLONURIDAE.
By R. J. Tinrtyarp, M.A., Sc.D: (Cantab.), D:Se. (Syd.), F-.R:S.,
AVNEZAnStls Hess. HGS RAES. (C.MeZ:s.

(Plate i; forty-five Text-figures. )
[Read 29th March, 1933.

Introduction.

Like South Africa and other countries subject to periods of drought,
Australia is on the whole extremely poor in Mayflies. This is clearly due to
the fact that the larvae, or nymphs, of most kinds of Mayflies require running
water in which to live. It might reasonably be expected, therefore, that those
parts of Australia where perennial, fast-running streams exist would prove an
exception to the general rule. The most elevated region in the whole of
Australia, and the one which possesses the most rapid rivers, is the Mount
Kosciusko range in the extreme south of New South Wales.

The Mayfly fauna of this region first attracted my attention in 1905. Mayflies,
however, are not like dragonflies, easily collected and worked out systematically.
Most of the adults have an extremely short life, and one would be extremely
fortunate if, during the course of a short visit to the mountain, one happened to
strike the exact time of year at which a particular adult Mayfly was on the
wing. Further, very few of the Australian species of Mayflies occur in large
numbers or fly in swarms, and I suspect that some of them are nocturnal in
their habits. Thus the collector of these insects will meet at the start with the
very annoying but unavoidable experience of collecting large numbers of larvae
or nymphs in the streams and finding himself unable, except at the inconvenience
of a prolonged stay, to determine the adult form, either by rearing the larvae
or by observing the actual emergence of the fly on the streams. To take an
example—the nymphs of the huge species of Coloburiscus described in this paper
were first discovered by me on Diggers’ Creek (5,000 ft. level) in January, 1905.
During a number of subsequent visits I endeavoured to rear or otherwise discover
the imago. Visits in late November, late December and early January all
indicated that this species most probably did not emerge until February. It
was not until 1929 that I was able to arrange a visit which lasted into the first
week of that month, when I discovered that the adult Mayfly emerged between
sunset and sunrise and almost immediately rose into the air and disappeared
into the bush far away. The following year I timed my visit a little later, and
secured a fine haul of subimagos. Not a single imago, however, was seen at any
time, and those which I possess were all reared by keeping the subimagos in
large tins, resting on reed-stems stuck into a layer of wet sphagnum moss. By

D

2 MAYFLIES OF THE MOUNT KOSCIUSKO REGION. i,

this method, it seemed at last that the problem of the giant Mayfly of Mt.
Kosciusko had been solved. However, a further visit was made in company
with Professor W. M. Wheeler early in December, 1931, at a time of year at
which I had never previously visited the mountain. Much to my surprise, I
found, in the same locality on Diggers’ Creek, and elsewhere, a number of
evidently full-grown nymphs of Coloburiscus, apparently indistinguishable from
those found in February. In a day or two, these began to emerge in small
numbers, when they were found to consist of an almost equally large but
entirely distinct species, with a differently coloured subimago.

In my book on Australian Insects (1926, p. 61), I have listed four families
of Mayflies as occurring in Australia, viz., Siphlonuridae, EHphemeridae, Lepto-
phlebiidae and Baétidae. This list has now to be modified in at least two
directions. Firstly, the minute Mayflies of the family Brachycercidae have been
recorded from Tasmania by Lestage (1930), and I am also able to state that they
occur in swarms on the Murrumbidgee River not far from Canberra. Secondly,
the record for the family Hphemeridae was based on two larvae discovered by
me in 1917 in the Fish River, N.S.W., but never described or reared. These
larvae were of the burrowing type, with large calliper-like projections in front
of the head. Unfortunately, the tube containing them was lost when I moved
to New Zealand; but I never had any doubt that they belonged to the Ephemeridae.
Quite recently, in dredging in the Murrumbidgee River in gravelly rapids and
under loose rocks, I rediscovered this same larva. On examination, I was
astonished to find that the large calliper-like processes are not part of the
mandibles at all, as they are in the Ephemeridae, but are frontal projections
designed to serve the same purpose, viz., to burrow down into the river-bed.
It is now clear that this larva does not belong to the Ephemeridae at all, but
is probably a highly specialized type developed from the Leptophlebiidae.

It is, of course, quite probable that true species of Ephemeridae occur in
Australia. If so, they have not yet been discovered. Likely localities in
which to search for them appear to me to be Western Australia and aiso the
Cape York Peninsula and the large, rapid rivers as far south as the Bellenden
Ker Range.

A further addition to the Australian Mayfly fauna may still be necessary,
owing to the discovery in the Cotter River, near Canberra, of a single specimen
of an entirely new type of Mayfly nymph which I am at present unable to place
in any recognized family.

The foregoing account will have made clear one important point, viz., that
it is much easier to discover new types of Mayfly nymphs than it is to determine
their corresponding imagos. Following the example of Needham and his school,
it is now quite usual, not only to found new genera on nymphal characters alone,
but also to describe new genera and species from nymphal forms, leaving it to
the effluxion of time to provide information about the imagos. A striking recent
example of the success of this method was the discovery and description of the
European genus Yorleya (Lestage, 1917). Much as one may dislike basing
generic and specific descriptions on larval forms, it seems to be the right method
of attack in studying a group like the Mayflies, where the life of the imago has
been reduced to an extremely short period and the whole struggle for existence
has been transferred to the larva. It is, indeed, absolutely true that, in most
cases, the Mayfly nymph provides much more reliable characters for classification
than does the imago. Lestage’s diagnosis of Torleya from the characters of its

ee)

BY R. J. TILLYARD,

nymph proved to be extraordinarily correct when, later on, the imago was
fortunately discovered.

A prejudice against the method of describing Mayfly nymphs without their
corresponding imagos has, I confess, hitherto prevented me from attempting
to write an account of the Mount Kosciusko species. Now, however, I have no
further excuse, for the imagos of all the more important species have at last
been determined.

The present paper will deal entirely with the Mayflies of the Mount Kosciusko
region from the summit down to about the 3,000 ft. level, the lower boundary
being taken along the Snowy and Thredbo Rivers to their junction at Waste
Point, not far from “The Creel’. The highest points at which Mayfly nymphs
have been taken are Lake Cootapatamba (6,600 ft.), Lake Albina (6,350 ft.)
and the Blue Lake (6,200 ft.). Specimens of adult Siphlonuridae have been taken
py me resting on rocks above snow, not far from the summit, at 7,000 ft. The
Mayfly fauna of these Alpine lakes is of very great interest, the dominant forms
being Siphlonuridae.

Only two families are so far known to occur on Mount Kosciusko, viz., the
Siphlonuridae and the Leptophlebiidae. Brachycercidae certainly occur lower
down along the Snowy River, round about Jindabyne, and probably Baétidae
also, so it may be possible to add these two families to the record later on. In
this part, I propose to deal only with the Siphlonuridae, which form the dominant
group of this elevated region and include some of the largest and finest Mayflies
in the whole world.

In attempting to classify the known Australian types of this family, one
is faced immediately with the fact that they are very closely allied to the New
Zealand forms of the same family, and that these latter forms are still in need
of revision, in spite of the excellent work recently carried out on them by
Captain J. S. Phillips (1930). Acting on a suggestion made to him by me
in litt., this author has removed Ameletus perscitus Eaton from the genus Ameletus
and made it the type of the genus Ameletopsis Phill., a proceeding fully justified.
This leaves two New Zealand species still within the genus Ameletus, viz., A.
ornatus Haton and A. flavitinctus Till. The type of the genus Ameletus, however,
is A. subnotatus Eaton from. Colorado, U.S.A. It has for, long appeared to me
unlikely that the New Zealand species belonged to the same genus as the American
type. Through the kindness of Dr. J. MacDunnough of the Division of Entomology,
Ottawa, Ont., Canada, I have recently received material in alcohol of both adults
and nymphs of this and other closely related North American genera. A study
of these reveals the fact that the New Zealand species are not congeneric with
them, as I long ago suspected.

It seems best, therefore, in defining the Australian genera, to bring the
New Zealand forms into the discussion and to construct keys which will include
both the Australian and New Zealand forms, both for adults and for nymphs.
It will then be clearly seen that there is only a single genus common to the
two countries, viz., Coloburiscus Eaton, which is the dominant and most widely
spread genus of the family in both, and that the other genera, though closely
related, are quite distinct from one another. A new generic name is required
for the New Zealand species at present placed in Ameletus Eaton; for these I
propose the name Nesameletus, n.g., fixing the genotype as Ameletus ornatus
Eaton. The other New Zealand genera are Oniscigaster McLach., Ameletopsis Phill.,
and Coloburiscus Eaton.

4 MAYFLIES OF THE MOUNT KOSCIUSKO REGION. i,

Family SIPHLONURIDAE.

Adults——Forewing with the tornus at from two-fifths to nearly one-half of
the wing-length, with vein CuA straight or nearly so, ending up just beyond
tornus and provided with a pectinate series of descending branches; CuP short
and curved concavely to CuA. Hindwing comparatively large, from one-third
to one-half as long as forewing, its venation complete, with triads present on
Rs and MA. Turban eyes never present in males. Subimago able to live in a
quiescent manner for a period ranging from two to four days.

Nymphs.—Free-living, non-burrowing types, with the mandibles not provided
with tusks; habits either free-swimming or else clinging to rocks in fast-running
streams; vegetarian or carnivorous. Gills usually simple, held either laterally
or dorsally. Tarsal claws usually smooth. Tail-filaments short to medium (not
longer than the body). Metamorphosis usually takes place while the nymph is at
rest on a rock, stone, reed-stem or on the bank of the stream just above water-level.

In dealing with the wing-venation of the adults (Text-figs. 1, 17, 28, 32, 43), I
have adopted the Revised Notation as set out in my most recent work on fossil
Mayflies (1932). In this notation, the two elements of the media are clearly
distinguished, viz., the anterior convex median, MA, and the posterior concave
median, MP, the latter corresponding with Comstock’s media M in most Orders.
The limits of the radial sector, Rs, are clearly defined if one remembers that the
base of this vein, in all recent Mayflies, has been secondarily captured by the
upwardly arching stem of MA. In order to bring the terminology of the cubitus
into line with that of the media, the anterior convex cubitus, hitherto called
Cu,, is now termed CuA, while the posterior concave cubitus, hitherto known as
Cu,, is termed CuP. The anal veins are termed A,, A,, A, and A,, respectively,
so as to allow of the use of the prefix “I” for intercalated sectors. The term
“triad” is used for a dichotomy of a main vein together with an included inter-
calated sector of opposite sign; this latter vein always takes the prefix “I” and
is named from the vein just anterior to it. The complete venational scheme
for the family can be studied in Text-figures 1, 17, 32.

Key to the Australian and New Zealand Genera.
Adults.

1. In forewing MP, and frequently also IMP are attached basally to CuA. Tarsal
claws alike, both sharp. Three well-developed tail-filaments in both sexes,
the appendix dorsalis being nearly as long as the cerci. (Hindwing with costa
onlyishehtlyaangcuilatedanears bases) see eee Ameletoides, n.g-

In forewing, MP, and IMP are normal, not attached to CuA. Tarsal claws dis-
similar, one sharp and one blunt. Appendix dorsalis either much shorter than

Cerciwopsolescents) on pentinely--abSentan sewers hee Oe eee eee 2

2. Tarsi apparently only four-segmented (the true basal segment of the tarsus being
fusedoywithwthesCibiaiy | Byers ceevekess cis eer ol ne eee ee oe Tasmanophlebia Till.

Tarsi- clearlysonve=SePmented © as cispociei dec heroves eit hem nee eee iehe oe EER eI ERO ee 3

3. Hind tarsus shorter than tibia. Hindwing with costa very strongly angulated near
IDA SER Ee ee tes icntep ey Me Reds Pipe out bation ihe coyotes uectoue evsitere onebepetene ote he one ree eee techs Coloburiscus Eaton,

Hind tarsus longer than tibia. Hindwing with costa either weakly angulated or
TOUNGEA! CAPE ASE wersire cies ols yros fee «aise sre goings) oo Tome ome ts al eran sac oikees: MOBS Sh Cee eae aE eee eS 4

4. Comparatively stout-bodied species, with or without lateral dilatations of the
abdominal segments. Appendix dorsalis present, but much shorter than the

Cole) Glo) lee ENE RP At sy i rice cit os at OAC CREE RE OL OME CET iin Ind OtS iosmaciioinrs tio o Oniscigaster MecLach.
More slender-bodied forms, with cylindrical abdominal segments. Appendix dorsalis
either obsolescentaormentinelyaibSent . re ca cieicnerclies iether iio nereienenere nei ie en none iene 5

5. Cross-veins very numerous and regularly spaced; pterostigma with branched vein-
lets forming a double series. Genital forceps of male with only three segments
A samaene: sv Me isrieltcesy 58 GRetane Home Re Maro ENCE Ones ciseaiia) sonetehte raters Ree eneee nen sah ey ake asta Ameletopsis Phill.

(Jt)

oT

BY R. J. TILLYARD.

Cross-veins much less numerous and irregularly spaced; pterostigma wlth a
single series of unbranched veinlets. Genital forceps of male four-segmented.
9010 WO.3 Oo DOO Od OOw Cho t0 OU O.C10'O FONT EC IDEOIOROTE IG ET One CREO ON Oy Ce OCHO ID Nesameletus, n.Z-

Nymphs.
Nymphs clinging to rocks in rapids. ‘Thorax strongly humped, very broad. Gills
deeply bifid, held vertically over abdomen. ................ Coloburiscus Baton,
Nymphs free-swimming, shrimp-like. Thorax not exceptionally broad, either not
him pedmonmoniveshohtlyason mGalll’Silanirellaten om... see eis sl cleus ciel ctaiele ise eieerenen enone 2
Nymphs dorso-ventrally flattened, the gills carried dorsally on the abdominal
SCLITICTVUS Mane Wt ateMetotan-naN Pencil chobenen ion ciicls) olaieaens eiieiici sr she) cifere/<hetelisiisisitelel elon stan sirsiieleleiepouarene 3
Nymphs slightly laterally flattened, the gills carried laterally ................ 4
Six or seven pairs of gills, on abdominal segments 1-6 or 1-7 respectively, all
lamellate and resting flatly upon the abdomen ........ Oniscigaster McLach.

Only four pairs of gills, on segments 1-4 respectively; of these, the first pair are
large and opaque, forming an operculum or protective covering for the other
MG ECM DD ALEStr tent hed cheucncheieiercieicnenccehensueys\cueuchanemetepven dieters oie wis vse, « Tasmanophlebia Till.
Nymphs carnivorous, with huge head and large eyes; incisors of mandible separate,
each subdivided into two sharp prongs; palpi with numerous small segments
MPa Welton ete clic hetche ket (sitelievis eke afer siaifetre. tes ache ue ane Ree rchersuchS shaaatoe race AMELELODSiSE re MIL:
Nymphs plankton feeders, with normal head; incisors of mandibles fused together

into a single blunt tooth; palpi normal, three-segmented .................. 5
Tarsal claws with fine serrations. underneath. Appendix dorsalis markedly shorter
ULL AIT EL GOL Claes Were AO EheP AOA sc, Mata cd lies sicohe Bacatawenelemaiclicie ote naieier dinate ets Nesameletus, n.g.-

Tarsal claws smooth. The three tail filaments of approximately equal length.
RPT eeu K oust eset Bene ones MSN SS ONS 2d cite ps lojre 75 ls cove tone) oe: enspepisyansyers aioe solebae reversye ays Ameletoides, n.g.

It will be seen that the nymphs fall into three well marked types, each of

which is represented in both countries, as follows:

(oN)

by

®
Type of Nymph. Genera.
Australia. New Zealand.

Slow, free-swimming type, dorso- Tasmanophlebia Till. Oniscigaster McLach.
ventrally flattened; gills held
dorsally flat above the
abdomen.

Fast, free-swimming type, later- Ameletoides, n.g. Nesameletus, n.g.
ally flattened; gills held Ameletopsis, n.g.
laterally.

Sedentary type, clinging to rocks Coloburiscus Eaton. Coloburiscus Baton.

in rapids, with huge, humped
thorax and gills bilobed and
held vertically above the
abdomen.

Thus it can be seen that, while Coloburiscus is represented in both countries
closely similar forms, Tasmanophlebia is the Australian representative of

Oniscigaster, and Ameletoides is the Australian representative of Nesameletus.
A single nymph of the Ameletopsis type has been captured in a tributary of the
Cotter River, near Canberra, F.C.T., but is not dealt with in this paper.

Genus AMELETOIDES, n.g. Text-figs. 1-11; Plate i, figs. 1-2.
Adult.—Forewing (fig. 1) with tornus at about two-fifths of wing-length;

MP, is secondarily attached to CuA; bullae present. Hindwing (fig. 1) distinctly
less than half as long as fore, its costa only slightly angulated near base. Venation
cf both wings delicate but well formed, the cross-vein system being quite as
strongly chitinized as the greater portion of the main veins. Forelegs (fig. 24)
of the male imago longer than the body, about as long as the forewing; those of
female imago shorter than the body. Tarsal claws (fig. 28) of all legs similar,
both sharply pointed, smooth. Tarsi with first segment shorter than the second;

6 MAYFLIES OF THE MOUNT KOSCIUSKO REGION. i,

the hind tarsi about equal in length to the hind tibiae. Three tail-filaments present,
the appendix dorsalis almost as long as the cerci.

Nymph (fig. 4).—Free-swimming, rather shrimp-like, somewhat laterally
flattened and with thorax slightly humped (machiloid type). Head small,
hypognathous, the eyes lateral, the antennae short; mandibles (fig. 6) with a
single large incisor; maxillary and labial palpi three-segmented. Abdomen taper-
ing posteriorly; seven pairs of gills (fig. 10), one pair on each of segments 1-7;
gills simple, lamellate, consisting of an oval portion, supported by a chitinized
rim, and an unsupported dorsal flange; tail-filaments (fig. 11) short, about half
as long as the body.

Genotype, Ameletoides lacus-albinae, n. sp.

Habitat.—Lake Albina, Mount Kosciusko Range, 6,350 ft.

This interesting genus, which appears to be maintaining the last stage of
a precarious existence in Lake Albina, to which the introduced trout has not yet
penetrated, is undoubtedly the Australian representative of the group which passes
at present under the generic name of Ameletus in New Zealand, but for which a
new generic name (Nesameletus, n.g.) is being proposed in this paper. It can,
however, be distinguished at once from the New Zealand forms by certain char-
acters both of the adult and the nymph. In the adult, the tarsal claws are
similar, both sharp; those of the New Zealand forms I find to be definitely
dissimilar. Both subimago and imago have the appendix dorsalis well preserved,
nearly as long as the cerci; in the New Zealand forms, the appendix dorsalis is
reduced to a mere remnant. In the nymph, the single incisor of the mandible
is very broad and truncated, and the prostheca is inserted almost in contact
with it; the three tail-filaments are of almost equal length.

AMELETOIDES LACUS-ALBINAE, hh. sp. Text-figs. 1-11.
Imago. Text-figs. 1-3.

3d (dried): total length (excluding tail-filaments) 14-0 mm.; abdomen 9-7 mm.;
forewing 14 mm.; hindwing 6 mm.; cerci 17 mm.; appendix dorsalis 15 mm.

Head (somewhat shrivelled) small, black above, mostly brownish below and
in front; eyes black with pale brownish rims; ocelli colourless, glassy, the median
one much smaller than the laterals; antennae blackish, set in pale brown toruli.

Thorax shining black above; sides partly black and partly brownish; sternites
brownish-black. Legs: forelegs 14 mm. long, femur dark brown with black knee,
tibia dark brown shading to black distally, tarsus black; femur about 4 mm.
long, tibia somewhat shorter, tarsal segments in descending order of length 3, 2,
4, 1, 5; middle and hind legs only about half as long, femora and tibiae pale
semi-transparent brownish, knees broadly black, tarsi shading to blackish; tarsal
segments in descending order of length, 1 = 2, 5, 3, 4; tarsal claws (fig. 2B) both
sharp, similar, inner margins broadly curved, with a sharply angulated flange
of delicate, transparent chitin.

Wings (fig. 1) brilliant, the membrane hyaline, the veins well-chitinized,
brownish, except in the anal areas, where they are weak and almost colourless.
A slight brownish tinge is present in the pterostigmatic area of forewing. Cross-
veins moderately numerous, very irregularly placed.

Forewing (fig. 1) with 11-14 pterostigmatic veinlets, usually single, but a few
may be connected by cross-bars; two bullae present, the anterior about half-way
along Sc, the posterior immediately below it at the origin of R,. The most
interesting venational feature of the forewing is the attachment of MP. to CuA

BY R. J. TILLYARD. 7

(fig. 1); in some specimens IMP is incomplete basally, in others (as in the one
figured) it is attached basally to MP,, and thus it appears at first sight almost
as if this triad belonged to CuA instead of to MP. Hindwing (fig. 1) with the
humeral angle well-formed, obtuse, slightly rounded; the costal space markedly
narrowed just before half-way; triad of MP arising close to base, very large
and deep.

Abdomen: segment 1 as broad as thorax, to which it is closely applied, black;
segments 2-9 strongly banded, basally pale brown, distally black; segment 10 pale
brownish. Genitalia as in figure 3, the penis pale brownish, large, with widely
diverging lobes; forceps-basis broad, pale brownish; forceps with four segments,
brownish basally darkening to blackish on last two segments; first segment
broad, short; second segment narrower, very long; third segment narrow,
cylindrical, about as long as first; fourth segment shorter than third, narrow,
cylindrical with rounded tip. Tail-filaments with about 50 segments, pale
brownish with blackish annulations at the joints.

© (dried): total length 14 mm.; forewing 16 mm.; hindwing 7 mm.; cerci
14 mm.; appendix dorsalis 12 mm. Differs from the male in the smaller eyes,
the thorax with rich brown markings above, the forelegs only 11 mm. long and
having the segments of the tarsus, in descending order of length, 1 = 2, 3, 5, 4;
forewings somewhat paler with a tinge of lemon-yellow in the pterostigmatic
area; abdomen somewhat broader basally and more tapering distally; subanal
plate trapezoidal, its free distal margin strongly emarginate.

Subimago. Plate i, figs. 1, 2.

The chief difference between subimago and imago in both sexes is the colora-
tion of the wings, which, instead of being hyaline, are dull and irregularly mottled
all over with fuscous streaks and blotches. In general, each crossvein is surrounded
by a narrow band of fuscous, on which it stands out as a pale, whitish line. In
certain parts of the wing, these dark bands tend to run together, forming
blotches; in the forewing, there is a blotch over the two bullae, a wider one
below the pterostigma, one across the triad of MA, another below the end of
A,, and there is more or less coalescence along the distal margin, in the costal
and subcostal spaces and below MP,; in the hindwing, most of the bands in the
distal half have coalesced and some of those in the basal half also. Generally
speaking, the mottling is darker in the male than in-the female, and there is
also more coalescence of individual bands. The forelegs of the male are consider-
ably shorter than in the imago, and the tail-filaments of both sexes are much
shorter also, the cerci being about 10 mm., the appendix dorsalis about 8 mm.
and both appearing rather shrivelled.

Nymph. Text-figs. 4-11.

Total length (excluding tail-filaments) 14-4 mm. Cerci 5 mm., the appendix
dorsalis almost equally long.

General form (fig. 4) somewhat shrimp-like, slightly compressed laterally,
with thorax slightly humped (machiloid type). Colour a uniform fuscous above,
shading to brownish below; gills greyish-white.

Head moderately large, hypognathous, with large, oval, black eyes and very
short antennae (1:5 mm.). Mouth-parts: Labrum (fig. 5) about twice as wide
as long, distal margin well rounded laterally, emarginate medially. Mandibles
(fig. 6) of markedly different shape, the right one being wider and more
triangular than the left; in both there is only a single large, turret-like incisor,

MAYFLIES OF THE MOUNT KOSCIUSKO REGION. i,

3

Fig. 1.—Ameletoides lacus-albinae, n.g. et sp. Venation of male imago.
Revised Notation: A, to A,, anal veins; CuA, anterior or first cubitus (convex) ;
CuP, posterior or second cubitus (concave); MA, anterior median (convex) ;
MP, posterior median (concave). The prefix “I’’ indicates an interpolated vein
forming the middle member of a triad; such a vein is of opposite sign to that
of the two veins enclosing it. Length of forewing, 14 mm. ®&

BY R. J. TILLYARD. 7]

very strongly and somewhat obliquely truncated distally; almost touching the
base of the incisor, on its inner side, is a short bristle representing the prostheca,
and, in the left mandible only, there is a longer bristle arising even closer to
the incisor and reaching beyond its apex; molar area of left mandible a rather
short combined grid and comb, the latter with about ten closely-set, slender
prongs, slightly curved at their tips; molar area of right mandible a much
longer but shallower grid only. Maxillae (fig. 7) with three-segmented palp,
the basal segment being the longest and broadest, the second slightly shorter,
much narrower, cylindrical, the distal segment very short and narrower still,
somewhat blunt at apex; galea and lacinia about of equal length, with their
apices close together, distal half of inner margin of lacinia with a fringe of stiff
hairs. Labium (fig. 8) with stout, three-segmented palps, the basal segments
longest and widest, bare except for two or three small, short setae near outer
margin, the second segment about two-thirds as long and not so wide, also
almost bare but with a small group of extremely delicate hairs near distal part
of inner margin, the distal segment short and fairly well rounded, with numerous
short setae more especially congregated on and near the distal margin; paraglossae
narrow, very elongated, reaching the middle of inner margin of distal segment
of palps, tips with delicate hairs and a few very short setae; glossae similar in
torm to the paraglossae but extending slightly beyond them, tips with a mass
of minute sensillae, two or three of which project as short, stout, blunt rods.
Hypopharynx extremely delicate and almost colourless, broad and rather truncate
distally, the paragnaths (superlinguae) well developed, covered with fine hairs
and carrying also a minute ridge of tiny teeth.

Thorax somewhat swollen and humped, the prothorax short and fairly
wide, convex above, the mesothorax not quite as long as broad, the wing-sheaths
broad and ending above the abdomen between the first pair of gills, the meta-
thorax short and for the most part hidden. Legs (fig. 9) set fairly close together,
the coxae short and stout, the trochanters short but well developed, the femora
nearly as long as tibia and tarsus combined, broad and fairly flat; tibia partly
divided into two by an oblique groove; tarsus slightly shorter than tibia, the
claw (fig. 2B) smooth, well pointed, with a row of five or six minute dots or
tubercles about middle of distal part.

Abdomen tapering markedly towards the posterior end, the segments becoming
longer from 1 to 4, then approximately equal in length from 4 to 9, 10 very short;
there is no dorsal crest or carina; posterior lateral angles of segments 3-9
inclusive end in a small, sharp spine; posterior margins of segments 4-9 inclusive
are finely crenulated. Gills (fig. 10), seven pairs, one pair on each of segments
1-7 inclusive, increasing in size from the first to fifth pairs, then decreasing; the
first pair are the smallest. Hach gill is inserted on the postero-lateral margin
just dorsally from the actual angle; it consists of a single broadly oval lamella
having its ventral margin strongly chitinized and its dorsal margin membranous;

Fig. 2.—Ameletoides lacus-albinae, n.g. et sp. Legs of male imago. 2A —
Tibia and tarsus of foreleg; tb tibia, ts tarsus (x 8). 2B.—Tarsal claws (x 75).

Fig. 3.—Ameletoides lacus-albinae, n.g. et sp. Genitalia of male imago
(x 30); fo forceps-basis, fe forceps, pe penis. Ventral view.

Fig. 4.—Ameletoides lacus-albinae, n.g. et sp. Nymph, nearly full-grown,
lateral view (x 6).

Figs. 5-8.—Ameletoides lacus-albinae, n.g. et sp. Mouth-parts of nymph.
5.—Labrum (x 30); 6.—Mandibles (x 30); ad adductor muscle, in incisor,
mo molar area, pr prostheca; 7.—Maxilla (x 30): 8.—Labium (x 30).

10 MAYFLIES OF THE MOUNT KOSCIUSKO REGION. i,

between the two runs a strong chitinous support, slightly curved and nearer
the dorsal than the ventral edge; the gill thus appears to consist of a strongly
supported ventral part between the stiff ventral margin and this support, with a
free dorsal flange readily movable in a current of water; the ventral margin is
finely toothed along its distal third. There is a single stout gill-trachea which
branches in all directions in the gill, but has its finest branches in the dorsal
fiange. The colour of the gills is milk-white shading to pale grey. Cerci (fig. 11)
stout, with short, annular segments and carrying a strong fringe of hairs on
the inner margin; appendix dorsalis similar, almost as long, and carrying a
fringe of hairs on either side.

Types.—Holotype male imago, allotype female imago, type male subimago
and type female subimago all taken at Lake Albina, Mount Kosciusko, 6,350 ft.,
the first three on 2nd Feb., 1929, the last-named on 1st Feb., 1930. Type nymph
and nymphal exuviae taken on rocks in the lake, 1st Feb., 1930. All the above
in the Tillyard Collection, together with a paratype series of six imagos and two
subimagos, pinned, and several in alcohol; also a series of nine slides with
mounts of the most important imaginal and nymphal structures, from which
the figures here given have been drawn.

This remarkable insect occurs, as far as is known, only on Lake Albina. I
have searched for its nymph in the other lakes and streams on Mount Kosciusko,
but have failed to find it there. The nymphs are confined to the waters of the
lake itself and of two or three very short streams which enter it from the
surrounding mountains. They may be seen either resting on the sandy bed, or

SS

TL)
Ss
DY

S
SS
\y
S
\

te

15 16

Figs. 9-11.—Ameletoides lacus-albinae, n.g. et sp. Parts of nymph. 9.—
Hind leg (x 16); fm femur, tb tibia, ts tarsus; 10.—Fifth gill (x 30); 11.—End
of abdomen and tail-filaments (x 12).

Figs. 12-16.—Details of Ameletus Eaton s. str. for comparison with
Ameletoides, n.g., and Nesameletus, n.g. 12.—Ameletuws velox Dodds (North
America) ; tarsal claws of imago (x 75); 13.—Ameletus ludens Needham (North
America); labrum of nymph (x 30); 14.—Ameletus velox Dodds; mandibles of
nymph (x 30); 15.—Ameletus ludens Needham; maxilla of nymph (x 30);
16.—Ameletus velox Dodds; anterior part of maxilla, to show plankton-rake
(x 30). Lettering as in fig. 6, except pl plankton-rake.

BY fk. .J. DILUYARD, 11

on rocks and stones. When approached with a net, they swim off quickly with
a darting motion. They are best caught by using two nets, keeping one fixed,
and gradually moving the other towards it, disturbing the sand and stones in
the way; then the nymphs resting on those stones dart forward into the
stationary net. The gills are either held stationary, or sometimes vibrate quite
rapidly. Nymphal exuviae are not found floating on the water, but resting on
the sides of rocks in the lake, an inch or two above water-level. Adults have
always proved very scarce during my visits to the lake; occasionally males may
be seen dancing in the air, but they appear mostly to prefer to rest on the shady
side of large vertical rocks, where they are very watchful and not easy to catch.
I have also taken specimens stuck on the snow which lies all the year round
near the summit of the mountain, at about 7,000 ft.

There can be no doubt that this is the single remaining Australian representa-
tive of the group represented by the New Zealand genus Nesameletus, n.g.
(previously Ameletus spp.). Both imagos and nymphs are closely similar in the
two genera, and the mottled colouring of the subimagos is much the same in
both. Only a few small but important characters serve to separate the two
genera, which are really more closely related to one another than either of them
is to Ameletus, s. str., a Holarctic genus.

In order to distinguish clearly the characters of these related forms, I
propose to include in this paper a discussion of the New Zealand genus.

Genus NESAMELETUS, 0.2.

This genus has been very fully defined by Phillips (1930, p. 311) under the
name “Genus AMELETUS (New Zealand Type)’. Some of the characters used
by him appear to be not of generic value, notably the presence of a median dark
band on the femur. I also think that Phillips is in error in defining the
claws as “alike, narrow and hooked in each tarsus”. This character also puzzled
Eaton (1885, 1899), for he first of all placed the New Zealand species ornatus
in Chirotonetes (claws alike), but later in Ameletus (claws dissimilar). The
truth appears to me to be that the claws are distinctly dissimilar, but that the
broadened claw is not as blunt as it is in the American and European species
of the genus Ameleius.

The genus may be defined as being very close to Ameletoides, n.g., from which
it differs only in the following points:

Imago.—Appendix dorsalis greatly reduced, 1 to 3 mm. long only; tarsal claws
distinctly dissimilar, one very sharp, the other broad and more or less blunt, but
not as markedly so as in Ameletus, s. str., or other genera of this family exclusive
of Isonychia. Forewing with the triad of MP normal; hindwing markedly smaller
than in Ameletoides, n.g.

Nymph.—In the mandibles, the large single incisor tooth is similar to that in
Ameletoides, n.g., though not quite so truncated; but the prostheca is further
removed from the incisor and consists of a short bristle accompanied by an
inwardly-directed brush or feathered bristle more than twice as long. The other
mouth-parts similar to those of Ameletoides, n.g., except that the lacinia of the
maxilla is somewhat shorter and blunter. The tibia is not subdivided by an
oblique groove; the tarsal claws are finely serrated on the inner margin. The
gills are similar to those of Ameletoides, n.g. The appendix dorsalis is markedly
shorter than the cerci.

Genotype, Ameletus ornatus Eaton, from New Zealand.

12 MAYFLIES OF THE MOUNT KOSCIUSKO REGION. 1,

The genus contains a second species, Nesameletus flavitinctus (Till.), also
from New Zealand.

A study of the genotype of the original genus Ameletus Eaton, s. lat. (A.
subnotatus Eaton), and other closely related North American species (figs. 12-16)
shows, without a shadow of doubt, that the Holarctic and New Zealand forms
are not congeneric. The imagos of Ameletus, s. str., are comparatively small
insects with extremely weak venation (hence the generic name); the main veins
are more or less transparent, the cross-veins practically invisible. The claws are
markedly dissimilar, one being very broad but sharply pointed, the other narrow
but extremely blunt and rounded at apex (fig. 12). The nymphs have somewhat
the same appearance as those of Nesameletus, n.g., and Ameletoides, n.g., but differ
in some very important characters. The labrum (fig. 13) is fully as long as
wide; the mandibles (fig. 14) are not triangular, but have the incisor and molar
areas sharply sepatated, making them very irregular in form; there are two
very slender incisors instead of one large stout one; the maxillae (fig. 15) are
of an entirely different form, with very slender palp and a broad inner lobe
(combined galea and lacinia) widened out distally and carrying along its distal
margin a “plankton-rake” (fig. 16, pl). In the North American species which I
have examined, the gills are oval without any very clear tracheation, and the
chitinous longitudinal support is missing.

Genus TASMANOPHLEBIA Tillyard. Figs. 17-30; Plate i, figs. 3-6.
Tasmanophlebia Tillyard, 1921, pp. 409-412, figs. 1 (male genitalia), 2 (venation) ;

1926, p. 62.

This interesting genus, which is the Australian representative of the New
Zealand genus Oniscigaster, was proposed by me for the reception of a single
Tasmanian species, 7. lacustris Till., of which only the adults are known. Since
then two species have been discovered on Mt. Kosciusko, and both adults and
nymphs are now known. Nymphs of a third species also occur in the Upper Cotter
River, F.C.T., but the adults have not yet been found. A study of the new
material indicates that the original definition of the genus requires some slight
emendation, and that excellent nymphal characters are also available for dis-
tinguishing it further from Oniscigaster and other Siphlonuridae.

Adult—Forewing with tornus at one-half of wing-length or even somewhat
beyond; triad of MP normal, arising very close to the origin of that vein; bullae
faintly indicated or absent. Hindwing about half as long as fore, or somewhat
more, and very nearly as wide, the humeral angle well-developed, obtuse; triad of
MP very small, distally placed. Venation strongly chitinized, inclusive of the
anal veins; cross-veins abundant and fairly regularly placed. Forelegs of male
imago nearly as long as forewing; those of female imago only half as long as
forewing, or less. Tarsi of all legs, except forelegs of male, at first sight
apparently only four-segmented, owing to the basal segment being more or less
fused with the tibia; this fused first segment about as long as the tibia, and longer
than any of the others. Tarsal claws of all legs strongly dissimilar, one sharply
hooked and one broad and blunt, with rounded apex. Abdomen narrowly
eylindrical, tapering posteriorly, without lateral dilatations on any of the segments.
Only two tail-filaments present, the cerci longer than the abdomen, the appendix
dorsalis reduced to a minute remnant.

Nymph.—Free-swimming, dorso-ventrally flattened, the thorax not humped;
abdominal segments with median dorsal crest and lateral flanges having strongly

BY R. J. TILLYARD. 3

projecting posterior angles. Head small, hypognathous, with laterally placed eyes
and short antennae; mandibles with two sharp incisors; maxillary and labial
palpi three-segmented; hypopharynx bilobed. Abdomen tapering posteriorly; only
four pairs of gills, carried dorsally upon the abdomen, one pair on each of
segments 1—4, the first a pair of strongly chitinized lamellae forming an operculum
to the other three, which fit under it owing to the shortening of the first four
abdominal segments; tail-filaments somewhat more than half as long as the
abdomen, the appendix dorsalis somewhat shorter and much less strongly built
than the cerci.

Genotype, Tasmanophlebia lacustris Till. (Lake Lilla, 3,200 ft., Cradle
Mountain, Tasmania).

This remarkable genus appears to be well represented in the lakes and streams
at high altitudes, both on the mainland and in Tasmania. In some respect it is
the most archaic type of Mayfly still existing, notably in the large size of the
hindwing. It is fairly closely related to the New Zealand genus Oniscigaster
MeLach., from which it is to be distinguished by the narrow, cylindrical abdomen
of the adults, without any lateral dilatations of the segments, by the generally
smaller size and by the peculiar specialization of the gills in the nymph. Nymphs
of Oniscigaster resemble those of Tasmanophlebia fairly closely in general form,
and, like them, possess a dorsal crest and lateral flanges on the abdominal segments;
but there are seven pairs of gills instead of four, and each gill consists of a fairly
hard plate of broadly oval shape, carried dorsally, flat upon the abdomen.

Key to the Species of the Genus Tasmanophlebia Till.

Imagos.
i, SPaGes OF Small gis ieee nba AMY wie, sodmoondanoadeonbadoognooUeUMUbeoS 2
Species of larger size, forewing 13-16 mm. ................ T. lacus-coerulei, n. sp.
2. Body dark brown; lobes of penis in male with terminal seta, forceps with four
SCPUMENUS Meeker se craqetart nets eneksbsger cee AVeU See osm alee srialic: o ese-c tel ae Mieie qhieeus T. lacustris Till.

Body black; lobes of penis without seta, forceps with only three segments ........
5 O56 Gl BIOs Oey OPER ORO EME G. 0 aD RISER ETERS sath oho, GLEE ae Pe CIE ES nvgnescens, Nl. Sp:

Subimagos.
1. Species of larger size (forewing 13-16 mm.) with the wings bicolorous; a narrow
triangular strip along distal margin of forewing, bounded by an oblique line
from middle of pterostigma to tornus, markedly darker than rest of wing;

distal margin of hindwing also darker than the rest .... J. lacus-coerulei, n. sp.

Species of smaller size (forewing 10-12 mm.) with the wings unicolorous or

TIC As iy dae S OMT eATSP NCR Cees Coto peta soe oR eseten ketene ey ae Roney ch cna Wwouereh oi cVelleWer cner a. cues woMtenaie caatpeileis 2

2. Body brown; wings greyish with a touch of brown at bases .... TJ. lacustris Till.

Body blackish; wings also appearing black in life when folded, dark grey when

SWRA GM OU Came ere te ches cacrchey oe ed oes sce ueeM a ore) cued chore cetwshece T. nigrescens, Nn. sp.
Nymphs.

1. Dorsal carina with large tooth-like projections on segments 1-5, not sharply pointed,

those of segments 3-5 subequal and larger than those of segments 1-2 ......
BE eee etar He neL ete chew omer ces kemeteirst ction oie feine arms eustioustiouepe Cou auaet ieteucysca scien skene T. nigrescens, n. Sp.
Dorsal carina with large tooth-like projections on segments 1-2, sharply pointed;
segment 3 with a much less elevated projection, also sharply pointed; segments
4-7 with flatter, posteriorly directed teeth progressively diminishing in size,
Woewe (Ope Keener Tt WAY Rael Soogoucnandoubpoapuoo uote T. lacus-coerulei, n. sp.

TASMANOPHLEBIA LACUS-COERULEI, n. Sp. Figs. 17-27; Pl. i, figs. 3, 4.
Imago. Figs. 17-19.
3 (dried): total length (excluding tail-filaments) 14 mm.; abdomen 9-5 mm.;
forewing 13:5 mm.; hindwing 7 mm.; cerci 20 mm.; appendix dorsalis 0-3 mm.

14

MAYFLIES OF THE MOUNT KOSCIUSKO REGION. i,

it
PT AE GIG ED

sm

J gee SS.

BY R. J. TILLYARD. 15

Head small, dark brown; eyes (collapsed) black with reddish-brown rims;
ocelli shining black; antennae dark brown, short, surrounded by an area of lighter
brown.

Thorax deep shining brown above, inclining to black; sides with varying
shades of brown; sternites pale brown; above the middle coxae and around the
bases of the wings are patches of bright orange-red. Legs: forelegs 13 mm. long,
blackish-brown; femur 3:5 mm., tibia 2-2 mm., tarsal segments in descending order
of length 1, 2 = 3, 4, 5; middle and hind legs much shorter, about 7 mm., rather
pale brownish in colour; the amount of fusion between tibia and metatarsus some-
what variable, tarsal segments in descending order of length 1, 2 = 3, 5, 4; tarsal
claws (fig. 18c) markedly dissimilar, one very sharply hooked and one broad and
blunt with well-rounded apex.

Wings brilliant, the membrane hyaline, the veins well-chitinized, including
those of the anal area, and mostly brown in colour. First and second axillaries
of forewing greyish with dark centres; third axillary, costal brace vein and
basal portions of Sc and R up to brace, orange-brown. All costal and subcostal
cross veins suffused with brown pigment; pterostigma right to apex suffused
with brown from costa to radius and carrying numerous cross-veins linked in
the middle portion into a double series by cross-bars. A faint bulla indicated
about half-way along R.,, by a minute brown cloud, with sometimes another below
it on R,,;; hindwing usually entirely hyaline, but sometimes a tinge of yellow
on membrane at base of both wings. Hindwing with posterior margin slightly
concave or emarginate at end of CuA.

Abdomen: segment 1 as broad as thorax, to which it is closely applied, very
dark brown; segments 2-9 medium brown with a dark brown pattern; on segments
2-4, a broad longitudinal dark band on dorsum, not clearly defined; on segments
5-9, two narrow longitudinal dark bands, separated by a paler stripe along the
mid-dorsal line; segment 10 dark brown with paler rim; sides with indistinct dark
brown bands; underside paler brown with darker blotches, paired on each segment.
Genitalia as in Fig. 19, brown, the penis long, with lobes not prominent, close
together, evenly rounded at apices; forceps-basis broad, the forceps with three
segments only, the first being broad basally and very long, tapering distally, the
second much shorter, cylindrical, the third narrower and somewhat shorter than
the second, also cylindrical. Tail-filaments: cerci pale brown at extreme base, the
rest medium brown, cylindrical, with about 45 not very clearly marked segments;
appendix dorsalis a mere remnant, very short, with 4 to 5 short segments.

© (dried): total length 16 mm.; forewing 16 mm.; hindwing 8-4 mm.; cerci
19 mm.; appendix dorsalis 0:2 mm. Differs from the male in the smaller eyes, in

Fig. 17.—Tasmanophlebia lacus-coerulei, n. sp. Venation of male imago.
Revised Notation, as in fig. 1. Length of forewing 13:5 mm.

Fig. 18.—Tasmanophlebia lacws-coerulei, n. sp. Legs of male imago.
18A.—Tibia and tarsus of foreleg (x 8). 18B.—Tibia and tarsus of hindleg
(x 8). 18C.—Tarsal claws of foreleg (x 75). tb tibia, ts, metatarsus.

Fig. 19.—Tasmanophlebia lacws-coerulei, n. sp. Genitalia of male imago
(x 380). Lettering as in fig. 8. (Right forceps omitted.) Ventral view.

Fig. 20.—Tasmanophlebia lacus-coerulei, n. sp. Nymph, full-grown (x 4).

Figs. 21-26.—Tasmanophlebia lacus-coerulei, n. sp. Mouthparts and leg of
nymph. 21.—Labrum (x 30); 22.—Mandibles (x 30), ad adductor muscle,
im incisors, mo molar area, pr prostheca; 23.—Maxilla (x 30); 24.—Labium
(x 80); 25.—Hypopharynx and paragnaths (x 30); 26.—Hindleg (x 15), fm
distal end of femur, tb tibia, ts tarsus.

16 MAYFLIES OF THE MOUNT KOSCIUSKO REGION. i,

a slight tinge of olive on the thorax, in the shorter forelegs which are dark
brown on the femora shading to a rich medium brown on tarsi, and in the large
amount of faint pale yellowish suffusion on the wings; subanal plate bifid, with
a triangular excision separating pointed lobes.

Subimago. Pl. i, figs. 3, 4.

In both sexes, the subimago is of somewhat darker body coloration than the
imago; the forelegs of the male are only about half as long as the forewing. The
wings are bicolorous; in the forewing, most of the membrane is pale grey, but
the distal border is more darkly shaded, the dividing line running from about
the middle of the pterostigma to the tornus; in the hindwing, the distal border
is narrowly darkened all round. The effect is actually brought about only partly
by a slightly deeper tinting of the distal membrane; it is chiefly due to the
distal cross-veins being clouded with dark grey. In the forewing, the costal brace,
basal parts of Se and R up to the brace, and most of the axillaries are orange-
brown, while the same colour tinges the pterostigma; from brace to pterostigma
the costal and subcostal cross-veins are black. In the hindwing there is a
touch of orange-brown at the base. In the female, the dark distal bordering of
the wings is slightly wider and more pronounced than in the male.

Nymph. Figs. 20-27, 30a.

Total length (excluding tail-filaments) 19 mm. Cerci 7-5 mm.; appendix
dorsalis 6-7 mm.

General form resembling the nymphs of the genus Oniscigaster McLach., but
slightly narrower and more elongated; body dorso-ventrally flattened. Colour a
dull fuscous above, without any marked pattern, shading to brownish beneath;
the abdominal crests and flanges pale brownish, semitransparent.

Head small, somewhat transverse, with rather small eyes placed at the postero-
lateral angles; antennae very short. Mouth-parts: Labrum (fig. 21) more than
twice as wide as long, broadly rounded antero-laterally and slightly emarginate
medially. Mandibles (fig. 22) very stout, with two sharp incisors, each either
entire or slightly toothed distally; prostheca consisting of a strong bristle together
with a longer process carrying a brush of hairs distally; molar area of right
mandible very prominent, strongly angulate, provided with a large grid; that
of the left mandible differently shaped, with a large comb and a hard grinding-
plate; the comb carries about ten closely-set, slender prongs, slightly curved.
Maxillae (fig. 23) with three-segmented palp, the basal segment the longest and
broadest, subcylindrical, the second segment about two-thirds as long, somewhat
narrower, with an inner apical bristle, the distal segment shorter still, distinctly
narrower, cylindrical, with rather bluntly rounded apex carrying a number of
short setae; inner lobe (combined galea and lacinia) rather broad, with outer
margin strongly curved, distal portion of inner margin with abundant hairs and
a shorter set of hairs placed distally between the two margins. Labium (fig. 24)
with stout, three-segmented palps having their basal segments standing out trans-
versely, second segment somewhat shorter, broadening distally, third segment
short and fairly broad, well-rounded distally, hairy; paraglossae short, with outer
margins greatly curved, apex pointed and turned inwards, slightly hairy; glossae
slightly shorter than paraglossae, broad and fairly straight, very close together,
somewhat truncate distally. Hypopharynx (fig. 25) oval, slightly longer than wide,
deeply bifid medially; paragnaths (superlinguae) about the same length, broadened
distally and carrying a fringe of incurving hairs on distal margin.

BY R. J. TILLYARD. 17

Thorax strongly built but not humped, the prothorax short but much wider
than the head, the mesothorax about as long as wide, with a slight but definite
colour-pattern dorsally; the metathorax very short; forewing-sheaths reaching
back to overlap the bases of the gills. Legs (fig. 26) short, fairly close
together, coxae and trochanters short, femora fairly long and stout, tibia and
tarsus closely fused together with an oblique sutural line, the tibia markedly
shorter than the tarsus; tarsal claw long and sharply pointed, smooth.

Abdomen markedly dorso-ventrally flattened, tapering gradually from segment
3 to posterior end. Segment 1 broader than thorax, segments 2-3 somewhat
broader still, segment 4 slightly narrower than 3, the others decreasing markedly in
width. Segment 1 about one-fourth as long as wide, about as long as segment 4;
segments 2-3 very short. Lateral flanges present on segments 1-9, all sharply
pointed posteriorly, as shown in fig. 30a. Dorsal crest (fig. 304) with large
nodding teeth on segments 1-2, a less prominent nodding tooth on segment 3,
and with much flatter, posteriorly directed teeth diminishing in size from segments
4-7, that of segment 7 being very small. Gills (fig. 27), four pairs, on segments
1—4 inclusive, decreasing in size from first to fourth pair, and so inserted that the
large first pair practically entirely cover the other three pairs, to which they
form opercula; first pair (fig. 274) broadly oval, strongly chitinized lamellae, with
obsolescent tracheation; second pair (fig. 27B) similar in shape, but smaller, and
with better developed tracheation; third pair (fig. 27c) smaller and shorter, lobed
basally; fourth pair (fig. 27p) still smaller, lobed basally and having a large
overfold. When examined in situ on the nymph, the gills appear to consist of
four pairs lying dorsally upon the base of the abdomen and covered by the
opercula, the apparent extra pair being actually the overfolds of the fourth (fig.
20). Colour of operculum dark brownish, of the other gills dull greyish. Cerci
(fig. 20) stout, incurved distally, with short, annular segments carrying a strong
fringe of hairs on the inner margin; appendix dorsalis paler and less strongly
formed, somewhat shorter, with a fringe of hairs on either side.

Types.—Holotype male imago and type male subimago taken at Blue Lake,
Mt. Kosciusko (6,200 ft.) on 1st Feb., 1930; allotype female imago and type
female subimago in same locality on 28th Jan., 1929. Type nymph and nymphal
exuviae taken on rocks in the lake, 28th Jan., 1929. All the above in the Tillyard
Collection, together with a paratype series consisting of 28 imagos and 27 sub-
imagos, all pinned, and nearly all females, and several in alcohol; also a series
of fourteen slides with mounts of the most important imaginal and nymphal
structures, from which the figures here given have been drawn.

So far, this very striking Mayfly has only been found in the Blue Lake,
where the nymph may be seen resting on the large rocks and boulders which
occur in it, or sometimes on the pebbly bottom. They are by no means easy to
catch, as they readily slip away from the advancing net; however, when swimming
freely, they are not at all fast. When about to emerge, they climb just out of
the water on the steep rocks and boulders of the foreshore of the lake and transform
there. Subimagos may be captured with comparative ease while resting on these
boulders, while imagos may be found either drifting with the wind near the
lake, or sometimes dancing in groups of two or three in the bright sunshine.
Specimens have also been found stuck in the snow banks near the summit, at
about 7,000 ft.

18 MAYFLIES OF THE MOUNT KOSCIUSKO REGION. i,

TASMANOPHLEBIA NIGRESCENS, 0. Sp. Figs. 28, 29, 308; Pl. i, figs. 5, 6.
Imago. Figs. 28, 29.
6 (dried): total length (excluding tail-filaments) 12 mm.; abdomen 8 mm.;
hindwing 4:7 mm.; cerci long, broken; appendix dorsalis

forewing 10:5 mm.;
0:15 mm.

Fig. 27.—Tasmanophlebia lacus-coerulei, n. sp. Gills of nymph. A, first
gill or operculum; B, second gill; C, third gill; D, fourth gill, showing over-
Lola yay Aula sl'5s

Fig. 28.—Tasmanophlebia nigrescens, n. sp.
Length 4:7 mm. Revised Notation. Venation as in fig. 1.

Fig. 29.—Tasmanophlebia nigrescens, n. sp. Genitalia of male imago (x 30).
fb forceps-basis, fe forceps, pe penis. (Right forceps omitted.) Ventral view.

Fig. 30.—Dorsal crests of nymphs of Tasmanophlebia. A.—T. lacus-coerulei,
B.—T. nigrescens, n. sp. 1-7, abdominal segments.

Hindwing of female imago.

n. sp.

BY R. J. TILLYARD. 19

Head small, blackish; eyes (collapsed) blackish; ocelli pale brownish;
antennae brown.

Thorax shining black, with a slight touch of rich dark brown above. Legs
pale brownish, darker at the joints; forelegs 9 mm. long, tarsal segments in
descending order of length 1, 2 = 3, 4, 5; middle and hind legs much shorter,
about 3-5 mm., tarsal segments in descending order 1, 5, 2, 3, 4; tarsal claws
markedly dissimilar, blackish.

Wings mostly hyaline, not brilliant, all the veins dark brown, well chitinized,
including the anals of both wings. In the forewing, the pterostigmatic area
carries numerous cross-veins, many of which are linked together by cross-bars to
form an irregular double series; this area is more or less deeply tinted with
transparent brownish; in one specimen, a rich brown tinge spreads basad along
costal and subcostal spaces and obliquely across the base of the wing to form a
brown cloud covering most of the anal area; third axillary orange-brown. MHind-
wing (fig. 28) hyaline, either tinged with brown near base, or with the brown
tint more widely spread and reaching well along costa. Hindwing with posterior
margin only very slightly emarginate at end of CuA.

Abdomen: segment 1 as broad as thorax, to which it is closely applied, black;
rest of abdomen tapering posteriorly, blackish, without any definite pattern.
Genitalia as in fig. 29, black, the penis moderately long, with lobes quite distinct
and well rounded at apices; forceps-basis broad, the forceps with three segments
only, of which the first is much the longest, the third shorter and narrower than
the second. Tail-filaments: cerci pale brown with dark joints; appendix dorsalis
a minute remnant.

@ (dried): measurements of body and wings as in male; cerci 14 mm.;
appendix dorsalis 0:25 mm. Differs from the male only in the small eyes, the
much shorter forelegs, 3-5 mm. long, the stouter abdomen, and the wings entirely
hyaline except for the brown tinge on the pterostigma and the orange-brown of the
third axillary; subanal plate slightly excised medially.

Subimago. Pl. i, figs. 5, 6.

In both sexes, the subimago differs in life from the imago in having the wings,
when folded together, appearing black. In the dried specimens, the wings are
shaded deep grey, the membrane actually being pale grey and the veins clouded
with dark grey. Viewed as a whole, the forewing shows a slight pattern suggest-
ing an indistinct lunule, slightly paler than the rest, crossing the wing so that
its outer and more distinct convex margin stretches from the beginning of the
pterostigma across to the tornus. Also the basal portion of the wing is somewhat
paler than the rest.

Nymph. Fig. 308.

Total length (excluding tail-fillaments) 14 to 16 mm. Cerci 5 to 6 mm:;
appendix dorsalis slightly shorter.

General form the same as that of the previous species; colour-pattern a
slightly mottled sandy brownish, with darker mid-longitudinal stripe along
abdomen; this pattern gives some degree of protective colouring to the nymph
as it rests on the sandy bottom of the stream. The mouth-parts differ only very
slightly from those of the previous species. The principal difference between
the two nymphs lies in the form of the dorsal crest (fig. 30B); in 7. nigrescens,
n. sp., the first five segments possess large nodding teeth, not very sharp, while
segments 6-7 are devoid of armature. Gills much as in the previous species,

20 MAYFLIES OF THE MOUNT KOSCIUSKO REGION. i,

but not quite as broad; the opercula (first pair) more oval than in the previous
species.

Types.—Holotype male imago taken at Spencer’s Creek, Mount Kosciusko
(5,700 ft.), 29th Jan., 1930; allotype female imago, type male subimago and type
female subimago in same locality, 31st Jan., 1929. Type nymph and nymphal
exuviae taken in same locality, 3lst Jan., 1929. All the above in the Tillyard
Collection, together with one paratype male imago, about fifty paratype female
imagos and subimagos, and numerous nymphs and exuviae; also a series of five
slides with mounts of the more important imaginal and nymphal structures,
from which the figures have been drawn.

This species comes fairly close to the genotype, 7’. lacustris Till., from Lake
Lilla, Cradle Mountain, Tasmania. It is about the same size, and appears to have
very similar habits. It is, however, easily distinguished in life by the general
black appearance of the subimagos; the male genitalia of the imagos are also
very distinct and the hindwings are of somewhat different shape. The species
occurs resting on the reeds bordering Spencer’s Creek along the stretch just
above the main road to the summit, not far from the new Bett’s Camp. The
creek here is sluggish and widens out in places, with reedy borders and very
little movement of the water, so that the conditions approximate to those found
along the sides of Lake Lilla. The nymphs can be seen resting on the sandy
bottom of the creek, though their colouring is partly protective. They are very
easily caught with a net. When emerging, they climb a few inches out of the
water up a reed stem, and transform there. The subimago crawls up the stem
and rests there, a conspicuous object. A very curious fact is that, out of more
than fifty subimagos collected, only three were males. When kept on reeds in
glass tubes, the subimagos transform on the second or third day, but many
fail to effect the transformation without damage to the wings. Trout eat this
species avidly; possibly the male subimagos fall victims to the trout through
flying across the water more readily than the females.

Genus CoLosuriscus Haton. Figs. 31-45; Pl. i, figs. 7-10.

Coloburus Eaton, 1868, Ent. Mo. Mag., v, p. 89 (genotype, Palingenia humeralis
Walker 1853); Eaton, 1871, Trans. Ent. Soc. London, p. 132; Eaton, 1883-88,
Revisional Monograph, pp. 201, 315.—Coloburiscus Eaton, 1883-88, Revisional
Monograph, p. 308; Eaton, 1899, Trans. Ent. Soc. London, p. 290; Hutton, 1898,
Trans. N.Z. Inst., xxxi, p. 217; Hudson, 1904, N.Z. Neuroptera, p. 35; Tillyard,
1926, Insects of Australia and N.Z., p. 62; Phillips, 1930, Revision N.Z.
Ephemeroptera, Part I, Trans. N.Z. Inst., \xi, p. 296; Phillips, 1931, Studies of
N.Z. Mayfly Nymphs, Trans. Ent. Soc. London, 1xxix, p. 414.

This is the only genus of Siphlonuridae common to Australia and New
Zealand, and is also undoubtedly the most highly specialized genus in the
tamily, found in those countries. The genotype, Coloburiscus humeralis (Walker),
is an extremely common species in most parts of New Zealand. The habits of
the nymph, which clings to rocks in rapids and also has a protective resemblance
to vegetable débris, have prevented this insect from falling too ready a prey
to the introduced trout. The North and South Island forms appear to me to
differ sufficiently for the former to deserve at least subspecific rank. In
Australia, so far, only one species has been described, viz., C. halewticus Haton,
from Victoria. Several other species are now known, but for the most part
the material available is not sufficient for description. There is a second very

BY R. J. TILLYARD. 21

distinct species in Victoria, another in the F.C.T., and a third in the Queensland
National Park, all from fairly high elevations. On Mount Kosciusko, two very
fine species occur, which are described in this paper.

Phillips (1929) indicates that in New Zealand the presence of nymphs of
Coloburiscus in a stream is a criterion of its excellence as a trout-stream. I
think the same holds good for the mainland of Australia; if so, the number of
first-class streams is much more limited.

Adult: Forewing with tornus at about two-fifths of wing-length from base;
triad of MP arising close to the origin of that vein, with MP, much curved near
its origin and approaching very close to CuA below it. Hindwing less than half
as long as forewing, the humeral angle strongly developed; triad of MP small,
distally placed. Venation strongly chitinized except at extreme base of wing
and on basal portions of anal veins; cross-veins abundant and fairly regularly
placed. Forelegs of male imago from two-thirds to nearly as long as the fore-
wing; those of female much shorter. Tarsi of all legs five-segmented, shorter
than the tibiae; tarsal claws of each leg strongly dissimilar, one sharply hooked
and one blunt; those of forelegs of male sometimes hypertrophied. Abdomen
tapering posteriorly, the first segment closely connected with the thorax; postero-
lateral angles of ninth tergite produced into spines. Only two tail-filaments
present, viz., the cerci, which are longer than the abdomen; the appendix
dorsalis reduced to a mere remnant.

Nymph.—Sedentary type, clinging to rocks in rapid waters. Body some-
what S-shaped in lateral view, the thorax huge and strongly humped, the
abdomen somewhat flattened dorso-ventrally, but without a dorsal crest. Head
of medium size, hypognathous, with fairly large eyes, laterally placed, and
antennae somewhat longer than the head; mandibles with two distinct incisors,
strongly projecting molar area and a large brush of long hair on the upper
surface; maxillary and labial palpi two-segmented, large; labrum, maxillae and
labium very hairy; hypopharynx entire. Thorax very large and convex; legs
rather short, spiny, the fore and middle femora also provided with long hairs;
tarsal claws simple. Abdomen somewhat dorso-ventrally flattened, tapering
posteriorly; gills carried upright dorsally upon the abdomen, one pair on each
of segments 1-7 inclusive; each gill is deeply bifid and strongly spinose; tail-
filaments three, the cerci well-developed, the appendix dorsalis variable.

Genotype, Coloburiscus humeralis (Walker), from New Zealand.

This peculiar genus is the most generally abundant Siphlonurid type
throughout New Zealand, while in Australia it is far more widely spread than
any of the other genera. In both countries it is far more likely to withstand
the attacks of the introduced trout than any other Siphlonurid type, owing to
the peculiar form and habits of the nymphs, which protect it to a large extent.

The genus is widely distributed on the mountain streams of south-eastern
Australia, wherever the flow of water is permanent and there are rocky rapids
suitable for the nymphs to inhabit. I have records of various species, still very
imperfectly known, from Victoria right up to south-eastern Queensland. There
is, however, so far no record from Tasmania, though one would expect the genus
to occur there. Mount Kosciusko may certainly be claimed as the headquarters
of the genus in Australia, since it is there that the nymphs are most abundant
and the species largest. Indeed, one of the two species described here is the
largest existing Mayfly known to me, with the sole exception of the giant Papuan
Mayfly (Plethogenesia papuana Eaton), an insect of a very different build and
appearance.

22 MAYFLIES OF THE MOUNT KOSCIUSKO REGION. 1,

In the following key I have introduced the New Zealand species C. humeralis
(Walker), in order to emphasize certain differences between the Australian
forms and the genotype. These differences, however, do not appear to render the
Australian forms deserving of even subgeneric rank.

Key to the Species of the Genus Coloburiscus Haton.

Imagos.
1. Medium-sized species, the forewing not more than 15 mm. .................. 2
Larger species, the forewing from 17 mm. up to 25 mm. ...........:......... 3
2. Forewing with a yellowish patch at base ............ C. humeralis (Walker), N.Z.
Morewine, without any, yellowish) patch 4.) sees oe _C. haleuticus Haton.

8. Very large species (forewing of male about 20 mm. long, of female up to 25 mm.)
with costal margin of forewing marked by a rich brown band ................-

RURAL A GIST ONO CHES HLCH- Hash A/G Ol cr UGS ER CII SHIN Geb ‘5 G'S oO OG Uae Gio olMiniOle'c C. giganteus, n. sp.
Species of somewhat smaller size (forewing of male 17 mm., that of female up to
22 mm.), the costal margin hyaline or only tinged with brown along the
PLerostiomar he 25: joes Ae. AO be eta een eirameeten ts. AU, Ueweh Stok one Ee C. munionga, n. sp.

Subimagos.
1. Medium-sized species, with a yellowish patch at base of forewing ..............
Sins BaC late dateuid waciouransanreclay te ede pe MES eH sia TS REN WATORSACLCRC MOIR IE De Helios ete catoans C. humeralis (Walker), N.Z.

lareer ispecies withoutwanyavellowishmpactchtnncajcinsse ror ie eee er oe Oe 2
2. Wings evenly suffused with opaque pale grey tint, the costal margin very faintly
to: moderately. ‘tinged swith: brown S23 seek oon ceae cee cin sole C. giganteus, n. Sp.

Wings mottled grey, the membrane very pale, the individual veins clouded with
medium grey; a small, irregular, clear space just beyond middle of forewing;
costal margin without any tinge of brown ................ C. munionga, n. sp.

(N.B.—The subimago of C. haleuticus Eaton is not known.)

Nymphs.
1. Appendix dorsalis less than half as long as the cerci; gills without any fibrils at
DASE) Ay PRR ech ats Wa aretcnemein ated ae leuelene ur oieien ace C. humeralis (Walker), N.Z.
Appendix dorsalis almost or quite as long as the cerci; gills with a large tuft of
fibrilsh Springing stron DAS hes simeiceckscticteleletere steer omteneredeueretcmspszo ac henene es renaee a peat 2
2. Very robust nymph of dark brown colour, with comparatively broad abdomen and
Heavily=SPINCAMlEESierreraiensner tence el cleo cision shenensMenet Meme orset cw C. giganteus, n. Sp.

Less robust nymph of a paler brown colour, with the legs less heavily spined
FP eth SOR OTD OTRO RECT E OO CRA OR CEG OM GOO como o ao Soe aol C. munionga, n. sp.
(N.B.—The nymph of C. haleuticus Eaton is not known.)
CoLOBURISCUS HALEUTICUS Eaton. Fig. 31.

Coloburus haleuticus Eaton, 1871, Trans. Ent. Soc. London, p. 133, Pl. vi,
figs. 7, 7a; Eaton, 1883-88, Revisional Monograph, p. 203, Pl. xviii, fig. 32c (penis),
Pl. xix, fig. 32 (forewing).

This species was described from a male imago taken in Victoria, “probably
near Melbourne (McCoy)”. There is little to go on in the description, which
is compressed into seven lines. The best characters appear to be that the fore-
wing is without any brown coloration along the costal margin and the penis is
of rather peculiar shape (fig. 31). Unfortunately the hindwing is not figured or
described.

I have not seen the type, which is probably in the Eaton Collection.

COLOBURISCUS GIGANTEUS, n. sp. Figs. 32-45; Pl. i, figs. 7, 8.
Imago. Figs. 32-34.
co (dried); total length (excluding tail-filaments) 16-5 mm.; abdomen 11-5
mm.; forewing 20 mm.; hindwing 8 mm.; cerci 23 mm.; appendix dorsalis 0:3 mm.
Head moderate, as wide as prothorax, blackish; eyes (collapsed) very large,
black with pale rims; ocelli pale amber, set in black; antennae very short, brown. |
Thorax medium shining brown above, slightly mottled, the mesonotum
projecting strongly backwards as a median cornute process; sides and underside

BY R. J. TILLYARD. 23

rich shining brown, with paler mottlings. Legs: forelegs 12-5 mm. long, brown,
the joints marked with black, the last tarsal segment mostly black; femur 3 mm.,
tibia 4-2 mm., tarsal segments in descending order of length 1 = 2, 3, 4 = 5; middle
and hind legs much shorter, the middle about 7 mm., the hind about 9 mm., both
pale, semi-transparent brownish, with black knees and touches of black at the
ends of the tarsal segments and along upper side of last tarsal segment and
claws; all the tarsal segments short, the complete tarsus considerably shorter
than the tibia; claws small, markedly dissimilar, the sharp claw with a delicate
tooth on inner margin; claws of foreleg hypertrophied into a broadly rounded,
dark process (fig. 334).

Wings brilliant, the membrane hyaline, the veins well-chitinized except at the
extreme base of the main veins of the forewing from Rs to CuP as well as the
anals. Forewing (fig. 32) with tornus at two-fifths of wing-length from base;
MP, arising close to base, either at or before the first cross-vein above it, and
curving so as to run much closer to CuA than to IMP. Pterostigma long, with a
complete double row of cellules, the connecting cross-bars forming very nearly a
complete line without any zigzagging. Costal and subcostal spaces entirely suffused
with brown, this colour deepening distally; third axillary brown, with a suffusion
of paler orange-brown posteriorly. Very small, faint bullae about half-way along
R.,3; and R,,;. Hindwing (fig. 32) entirely hyaline, fairly well rounded, the humeral
angle very prominent, the costa straight from this angle to beyond half-way;
costal cross-veins numerous, especially in basal half; anal veins moderately well
chitinized.

Abdomen: segments 1-2 very short, as broad as the thorax; segment 1 closely
affixed to thorax, medium brownish; segment 2 dark brown. Abdomen tapering
from segment 3 to end; each segment from 3 to 8 inclusive dark brown above,
with a pattern consisting of two round spots of paler brown placed anteriorly
and a slightly larger similar blotch in the middle line posteriorly; the two anterior
spots are clearly marked on segments 3-6, but become indistinct on qu and 8;
on the other hand, the posterior spot becomes more distinct on the ni6re distal
segments. Segment 9 dark brown, with ill-defined paler areas; its postero-lateral
angles carrying a sharp, backwardly projecting spine about two-thirds as long as
segment 10. Segment 10 itself mostly pale brown, darker on sides, and with two
small dark points placed antero-dorsally. Underside a much paler brown. Genitalia
as in fig. 34, mostly pale brownish, but distal part of forceps blackish; the penis
short, appearing in the dried specimen as a broadly rounded lobe not reaching
as far as the distal margins of the forceps-basis; after treatment with KOH, the
two short lobes can be seen, each subdivided into three incurving branches, of
which the most distal pair are crenulate on their outer margins; forceps-basis
divided into two halves, fairly broad, with rather prominent inner distal angles;
forceps three-segmented, the first segment much longer than the other two
combined, bent basally, then straight and cylindrical; the second segment little
more than one-third as long, narrower, also cylindrical; the third segment shorter
and narrower, very narrowly oval in shape. Tail-filaments: cerci very long, stout
basally, tapering to a fine thread distally, with from 60 to 70 segments (the distal
ones indistinct); colour rich brown, with blackish rings at the joints; appendix
dorsalis a minute, slender remnant consisting of 5-6 microscopic segments.

© (dried): total length 17 mm.; forewing 23-5 mm.; hindwing 9 mm.; cerci
20 mm.; appendix dorsalis 0-6 mm. Differs from the male in its considerably
larger size, smaller eyes, larger ocelli, shorter forelegs (about 10 mm. long)

24 MAYFLIES OF THE MOUNT KOSCIUSKO REGION. i,

Fig. 31.—Coloburiscus haleuticus Eaton. Lobes of penis. After Eaton.
Fig. 32.—Coloburiscus giganteus, n. sp. WVenation of male imago. Revised
Notation, as in fig. 1. Length of forewing 20 mm.

BY R. J. TILLYARD. 25

with normal claws, and much wider and more cylindrical abdomen. The broad
vertex between the eyes is brownish edged with a black line and, externally to
this, with a fairly wide rim of orange-brown around the eyes. The thoracic nota
are slightly paler than in the male. Except for segment 1, which is coloured like
the thorax, the abdomen is darker and without any marked pattern, except that
the sutures are pale. The postero-lateral spines of segment 9 are shorter and
wider, being only about half as long as segment 10. Subanal plate hollowed out
semicircularly on its distal margin, strongly bifid, with pointed lobes. Cerci
slenderer than in male, with only 35-40 segments.

Subimago. Pl. i, figs. 7, 8.

In both sexes, the brownish coloration of the imago is replaced by a much
duller greyish-brown, but the patterns remain the same; the forelegs of the male
are only half as long as the forewing. The wings are opaque, uniformly shaded in
pale greyish, without any trace of pattern, except that the costal and subcostal
spaces are more darkly shaded in greyish-brown. The cerci are much shorter
in the male, slightly shorter in the female. The postero-lateral spines in both sexes
reach to the end of segment 10 or slightly beyond.

The largest specimen of the series is the type female subimago, which expands
50 mm. or just on two inches.

Nymph. Figs. 35-48.

Total length (excluding tail-filaments) 17 to 20 mm. Cerci 10 to 11 mm.,
appendix dorsalis almost or quite as long as cerci.

General form rather stout and heavily built, the outline of the body viewed
laterally being somewhat S-shaped owing to the robust, strongly humped thorax
and the rather dorso-ventrally flattened abdomen, without dorsal crest. Colour
dark fuscous above, shading to medium brown on sides and paler brown beneath;
eyes black with pale rims; legs medium brown, the distal ends of the femora
and tibiae darker, spines dark brown; tail-filaments dark fuscous basally, shading
to medium brown distally. Gills medium brown with somewhat darker spines.
No definite colour-pattern present.

Head moderately large, strongly depressed, very convex dorsally and furnished
with a fringe of hairs anteriorly; eyes lateral, fairly large; antennae slender,
about 3 mm. long, brownish, composed of numerous short, ring-like segments.
Mouth parts: labrum (fig. 36) about twice as wide as long, very hairy all over,
broadly rounded antero-laterally and very slightly emarginate medially, with a
dark triangular patch in middle of anterior margin. Mandibles (fig. 37) very
stout, with two sharp incisors, each having its apex subdivided into three fine
teeth and each carrying, on its outer surface, a series of about thirty long,
slender hairs arising from the curved edge of a large shallow depression, and
almost covering the whole mandible, together with a smaller brush of shorter
hairs situated externally to the two incisors; right and left mandibles markedly

Fig. 33.—Coloburiscus giganteus, n. sp. Legs of male imago. A.—Tibia
and tarsus of foreleg, showing hypertrophied claws (x 8). B.—Tibia and
tarsus of hindleg (x 8). C.—Tarsal claws of hindleg (x 75). tb tibia,
ts tarsus. :

Fig. 34.—Coloburiscus giganteus, n. sp. A.—Genitalia of male imago (x 23),
ventral view, with right forceps omitted; fb forceps-basis, fe forceps, pe penis.
B.—Lobes of penis, further enlarged (x 45).

Fig. 35.—Coloburiscus giganteus, n. sp. Nymph, full-grown, lateral view
(x 4). ;

26 MAYFLIES OF THE MOUNT KOSCIUSKO REGION. i,

different in shape, the right one having the outer margin strongly curved, the
outer incisor strongly hooked, the inner incisor about as large as the outer but
not curved, the inner margin between this incisor and the molar area strongly
gouged out, and the molar area itself extremely prominent, wedge-shaped, with
well-developed grinding plates; no prostheca visible; the left mandible, on the
other hand, has the outer margin straighter, bluntly truncate before the outer
incisor, which is strongly hooked and considerably larger than the inner; the

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Figs. 36-41.—Coloburiscus giganteus, n. sp. Mouth-parts and end of
abdomen of nymph. 36.—Labrum (x 23); 37.—Mandibles (x 23), in incisors,
mo molar area, pr prostheca; 38.—Maxilla (x 23); 39.—Labium (x 28);
inside view of left half only; the other half is held parallel, with the thick
brushes of hairs in contact with one another; 40.—Hypopharynx and paragnaths
(x 23); 41.—End of abdomen and tail-filaments (part only), ventral view (x 8).

BY R. J. TILLYARD, 27

prostheca slender, longer than the incisor, with shorter pencil of inwardly directed
hairs; the margin between inner incisor and molar area perfectly straight, the
molar area itself at right angles to it, carrying a comb of closely-set, slender
prongs, fourteen or fifteen in number. Maxillae (fig. 38) with palps only two-
segmented, the basal segment rather short, smooth, the distal more than twice
as long, broader, sabre-shaped and very hairy; there is an indication from a slight
obliquely placed line near the base of this segment, bordered on one side with
hairs, that it actually consists of the two original distal segments fused together;
inner lobes (combined galea and lacinia) broad, cultriform, with a sharp tooth
at the inner distal angle; distal margin very hairy, inner margin with a close-set,
regular row of shorter hairs. Labium (fig. 39) with large, two-segmented palpi,
the basal segments smooth, long and fairly narrow, standing out transversely
or even directed somewhat backwards, the distal segments even longer, sabre-
shaped, very hairy, directed forwards; glossae and paraglossae overlapping on
each side, both broadly cultriform with bluntly rounded apices, both hairy, the
paraglossae more so than the glossae. Maxillae and labium held projecting
forwards in parallel vertical planes; the two halves of the labium (fig. 39) parallel
and close together, the angles of the palps projecting backwards between the
bases of the forelegs. Hypopharynx a small, weakly chitinized, broadly rounded
lobe, almost transparent, the paragnaths also rounded, not so broad, slightly
hairy. :
Thorax very stout, strongly convex above, the thoracic nota forming a kind
of hard carapace, almost black in colour, the pronotum being marked off by an
impressed transverse line, its length being about equal to that of the head viewed
from above; the broad, subtriangular forewing-sheaths are continuous with the
carapace and reach to between the second pair of gills. Legs (fig. 42) fairly large
and stout, approximately equal in length; foreleg with the femur broader and
shorter than in the other two, the tibia longer and more curved and swollen
near its base; the fore and middle femora carry a fringe of long hairs on a
little more than the basal half of the ventral margin, while the anterior surface
carries numerous small spines irregularly arranged, together with two transverse
halfi-rows of spines, of which the ventral half-row is placed distally from the
dorsal; the hind femora are spiny but not hairy; fore tibiae slightly curved and
widened basally, very spiny, with a long fringe of hairs on all but the extreme
base of the ventral margin; middle and hind tibiae very spiny, without hairs;
all tarsi short, about half as long as middle or hind tibia, smooth except for
a few minute setae at distal end; tarsal claws smooth, strongly hooked, fully
one-third as long as tarsi.

Abdomen stout, fairly broad, strongly tapering posteriorly, somewhat dorso-
ventrally flattened, without any dorsal crest; postero-lateral angles of segments
2-9 ending in a sharp spine, gradually increasing in size from 2 to 9; the larger
of these spines themselves carry a few small setae or spinules. Gills (fig. 43)
on segments 1 to 7 inclusive, one pair on each segment; all seven pairs large
and fairly equal in size, those of segments 1 and 7 a little smaller than the
others; each gill consists of a hardened chitinous process, deeply bifid, with two
sharply pointed prongs, both densely spined; from near the base of the gill, on
the posterior side, there arises a large tuft of delicate, greyish-white fibrils.
Tail-filaments (fig. 41) stout basally, tapering distally, straight, very slightly
hairy, held close together, diverging slightly distally; all three about the same
length.

28 MAYFLIES OF THE MOUNT KOSCIUSKO REGION. i,

Types.—Holotype male imago, allotype female imago, and type male and
female subimagos, all taken on Diggers’ Creek, Mount Kosciusko (5,000 ft.), on
30th Jan., 1930. Type nymph and nymphal exuviae taken under rocks in the
same locality, on the same day. All the above in the Tillyard Collection, together

Fig 42.—Coloburiscus giganteus, n. sp. Legs of nymph (x 12). A, foreleg;
B, middle leg; C, hindleg.

Fig. 43.—Coloburiscus giganteus, n. sp. Nymph, fifth gill (x 16).

Fig. 44.—Coloburiscus munionga, n. sp. Hindwing of male imago, Length
7-5 mm. Revised Notation. Venation as in fig. 1.

Fig. 45.—Coloburiscus munionga, n. sp. A.—Genitalia of male imago (x 23),
ventral view, with right forceps omitted, fb forceps-basis, fe forceps, pe penis.
B.—Lobes of penis, further enlarged (x 45).

BY R. J. TILLYARD. 29

with a series of eight paratype subimagos and sixteen paratype imagos, also
a number of nymphs, all from the same locality, 25th Jan. to 2nd Feb., 1930;
also a series of seven slides with mounts of the most important imaginal and
nymphal structures, from which the figures here given have been drawn.

This magnificent Mayfly is only to be taken towards the end of January or
beginning of February on Diggers’ Creek and other similarly rapid, rocky creeks
on Mount Kosciusko. The nymphs cluster in small colonies under rocks in the
swiftest parts of the stream. They were first found by me in 1905; but it was
not until 1930 that I saw the first subimago, resting on the edge of a rock, close
to the rushing water, at sundown. At the end of about half an hour or less
the subimago rises suddenly into the air at an angle of about forty-five degrees,
passing right out of sight, unless, as often happens, a bird seizes it. All my
subimagos were gathered from the rocks either near sunset or sunrise, and all
the imagos were reared from them, the average duration of subimaginal life
being two and a half days.

CoOLOBURISCUS MUNIONGA,* n. sp. Figs. 44, 45; Pl. i, figs. 9, 10.
Imago. Figs. 44, 45.

6 (dried): total length (excluding tail-fillaments) 16 mm.; abdomen 9 mm.;
forewing 17-5 mm.; hindwing 7-5 mm.; cerci about 20 mm.; appendix dorsalis
minute, slender, about 0:2 mm.

Head moderate, about as wide as prothorax, blackish; eyes (collapsed) large,
blackish; ocelli silvery, set on blackish prominences; antennae short, dark brown;
frons medium brown.

Thorax shining brown, the anterior and posterior processes of the mesonotum
pale brownish, with a dark median stripe between them, bordered on either side
by a pale stripe, and these externally by dark stripes; the two postero-lateral
bosses of the mesonotum orange-brown, shading to dark brown near the wings;
sides brown with paler spiracles; sterna rich brown. Legs: forelegs 12 mm.
long, the femur brown touched with blackish, the tibia blackish, the tarsus
brown with blackish joints; the tarsus about as long as the tibia; tarsal
Segments in descending order of length, 1 = 2, 3, 4 = 5; middle and hind legs
shorter, the hind about 9 mm., pale semi-transparent brownish, with darker
knees and touches of dark at ends of the tibiae and the tarsal segments; the
tarsus little more than half as long as the tibia; tarsal claws very small,
blackish, markedly dissimilar.

Wings brilliant, the membrane entirely hyaline except for a faint tinge of
brown on the pterostigma of the forewing. Forewing with tornus at about two-
fifths from base; venation similar to that of previous species, pterostigma with
a complete double row of cellules; third axillary narrowly dark brown, with a
greyish area posteriorly. Hindwing (fig. 44) much as in previous species, with
very prominent humeral angle and straight costa to beyond half-way, entirely
hyaline with dark axillaries.

Abdomen shaped much as in previous species, colour fairly dark brown, with
a pattern consisting of paler brown areas, arranged much as in previous species,
but larger and, on the whole, not quite as well defined; from segment 3 back-
wards there runs a fairly well-marked median longitudinal stripe of dark brown,
divided along the middle by a fine pale line; postero-lateral spines of segment 9
about as long as segment 10, very pale; segment 10 dark brown, bordered

* Munionga: the aboriginal name for the Mount Kosciusko Range.

30 MAYFLIES OF THE MOUNT KOSCIUSKO REGION. i,

posteriorly with pale brown; underside pale brownish, same colour as hind
legs. Genitalia as in fig. 45, mostly pale brownish, the penis somewhat darker;
in the dried specimens the penis is seen as a short, very stout process below
and between the cerci; after treatment with KOH, it is seen to have each lobe
subdivided into three processes, as in the previous species, but these processes
are very different in shape, two being broadly rounded and one rather slender,
while none of them is as incurved as those of the previous species; the outer
margins of the innermost pair of processes are barely crenulate and are minutely
punctate on the semi-transparent membrane just behind the strongly chitinized
outer border; forceps-basis divided into two halves, shaped much as in previous
species but not so large; forceps three-segmented, shaped much as in previous
species, but the short distal segment narrower and less rounded apically. Tail-
filaments: cerci with about 60 segments, brown ringed with blackish at joints,
not so stout as in previous species; appendix dorsalis a minute remnant with a
few indistinct segments carrying tiny setae.

@ (dried): total length 15 mm.; forewing 21 mm., hindwing 8 mm.; cerci
28 mm.; appendix dorsalis 0-2 mm. Differs from the male in its larger size,
smaller eyes, larger and much darker ocelli, shorter forelegs (9 mm. long) and
much wider and more cylindrical abdomen. The colouring of the vertex is
similar to that of the preceding species, but with a pale line dividing it longitu-
dinally. Colouring of thorax much as in the male. Abdomen with segments 1-2
medium brown, segments 3-10 very dark brown with the posterior margins of
segments 3-7 bordered by a transverse band of medium brown carrying, on
either side of the middle line, either two or three triangular patches of pale
brown with their apices projecting forwards; on segments 8-9 the pattern is
indistinct, while segment 10 is dark brown with a pale posterior edge; postero-
lateral spines of segment 9 shorter and wider than in male, about two-thirds as
long as segment 10, pale in colour; subanal plate deeply hollowed out. Cerci

stouter and longer than in the male, with more than 100 short segments, becoming
indistinct distally.

Subimago. Pl. i, figs. 9, 10.

In both sexes the brownish colouring of the imago is replaced by a duller
greyish-brown, but the colouring of the male subimago is closer to that of the
imago than in the female; the patterns in each sex remain very much the same.
The wings are mottled in pale and fairly dark grey, a marked pattern being
formed by the darkly shaded cross-veins on a membrane of very pale semi-opaque
greyish; each cross-vein is itself black, enclosed in a darkened area from
one-third to one-half as wide as long. In forewing, around the curved basal
portion of R,, several cross-veins have these dark areas fused together into a
single patch; below and somewhat anterior to this is a large pale patch of
irregular shape devoid of cross-veins and somewhat suggesting a lunule; hindwing
with the dark patches surrounding the cross-veins very narrow. Postero-lateral
spines of segment 9 of abdomen much as in imagos. Cerci 14 mm. in male,
23 mm. in female.

Nymph.

None of the nymphs which appear to belong to this species is full-fed, and
therefore I hesitate to offer characters for distinguishing it from those of the
previous species. The nymphs generally are of a much paler brown colour, some-
what less robust build, with slightly narrower thorax and more cylindrical

Proc. Linn. Soc. N.S.W., 19

99°
vo.

Subimagos of the family Siphlonuridae.

PLATE

ial

BY R. J. TILLYARD. 31

abdomen, the legs and gills less spiny. Mouth-parts closely resembling those
of previous species. How far the differences noted are due to immaturity I am
not able to say.

Types.—Holotype male imago taken at Diggers’ Creek, below waterfall (about
4,000 ft.), Mount Kosciusko, 12th Dec., 1931; allotype female imago and type
female subimago, both taken on Diggers’ Creek at 5,000 feet. on 11th Dec., 1931;
type male subimago taken on Spencer’s Creek (5,700 ft.), 31st Jan., 1929. Type
nymph (two-thirds grown) taken on rocks in Spencer’s Creek, 31st Jan., 1929.
ATl the above in the Tillyard Collection, together with four paratype imagos and
three paratype subimagos and a few nymphs; also a slide showing male genitalia
prepared from one paratype.

This species was originally discovered by me on Spencer’s Creek at the 5,700 ft.
level at the end of January, 1929, when two subimagos were taken and a single
imago was found damaged and floating down-stream. In December, 1931, when
visiting the mountain with Professor W. M. Wheeler and party, I was astonished
to find the same species appearing sparingly on Diggers’ Creek, not only in the
same places as those where C. giganteuws appears six weeks later, but also much
lower down. One of Professor Wheeler’s party secured two specimens not
far from the junction of the Snowy and Thredbo Rivers, at about 3,000 ft.

The species is easily distinguished from C. giganteus, n. sp., by its slenderer
build, smaller size, lack of the brown band along costa of forewing in the imago
and more especially by the strongly mottled wings of the subimago. The shape
of the penis of the male is characteristic also.

It should be noted that, while both the newly described species have penes
with each lobe subdivided into three processes, C. haleuticus Eaton has each lobe
with only two processes, and the New Zealand species, C. humeralis (Walker),
has the lobes undivided.

EXPLANATION OF PLATE I.
Subimagos of the family Siphlonuridae.
(All figures slightly more than natural size.)

1-2. Ameletoides lacus-albinae, n.g. et sp. Fig. 1, male; fig. 2, female.

3-4. Tasmanophlebia lacus-coerulei, n. sp. Fig. 3, male; fig. 4, female.
5-6. Tasmanophlebia nigrescens, n. sp. Fig. 5, male; fig. 6, female.
7-8. Coloburiscus giganteus, n. sp. Fig. 7, male; fig. 8, female.

9-10. Coloburiscus munionga, n. sp. Fig. 9, male; fig. 10, female.

References.

EATON, A. E., 18838-1888.—A Revisional Monograph of Recent Ephemeridae. Trans.
Linn. Soc. London (see pp. 199-230).

——__—__—.,, 1899.—An annotated list of the Ephemeridae of New Zealand. Trans. Ent. Soc.

London, pp. 285-2938.

Hupson, G. V., 1904.—New Zealand Neuroptera (see pp. 23-45).

LestaGE, J. A., 1917.—Contribution a l’étude des larves des Epheméres palaearctiques.
Ann. Biol. lacustre, viii, pp. 212-458. (Torleya, p. 366.)

, 1919.—Contribution 4 l’étude des Epheméres palaearctiques (Series 2). Ann.
Biol. lacustre, ix, pp. 79-182. (Siphlonuridae, pp. 162-174).

Puiuuies, J. S., 1929.—A report on the food of trout and other conditions affecting
their well-being in the Wellington District. Fisheries Bulletin No. 2, N.Z. Marine
Department.

,1930.—A Revision of New Zealand Ephemeroptera. Trans. N.Z. Inst., lvi,
pp. 271-334, plates 50-60.

,1931.—Studies of New Zealand Mayfly nymphs. Trans. Ent. Soc. London,
IXxix, pp. 399-422, plates XVi-xxili.

32 MAYFLIES OF THE MOUNT KOSCIUSKO REGION. 1.

TILLYARD, R. J., 1920.—Report on the Neuropteroid Insects of the Hot Springs Region,
N.Z., in relation to the problem of Trout-food. Proc. LINN. Soc. N.S.W., xlv, pp.
205-213 (also in N.Z. Journ. Sci. and Tech., 1921, iii, pp. 271-279).

,1921.—A New Genus and Species of Mayfly from Tasmania, belonging to
the Family Siphluridae. Proc. LINN. Soc. N.S.W., xlvi, pp. 409-412, pl. xxxiv.

, 1923a.—Descriptions of two new species of Mayflies from New Zealand. Trans.
N.Z. Inst., liv, pp. 226-230.

,1923b.—The Wing-Venation of the Order Plectoptera or Mayflies. Journ.
Linn. Soc. London, xxxv, pp. 143-162.

, 1926.—Insects of Australia and New Zealand (see pp. 57-64).

, 1932.—Kansas Permian Insects. Part 15. The Order Plectoptera. Amer.
Journ. Science, xxiii, pp. 97-134, 237-272 (Tasmanophlebia venation, figs. 9-12).

THE LIFE-HISTORY OF GREVILLEA ROBUSTA (Cunn.).
By P. BroucH, M.A., B.Sc., B.Sc.Agr., Lecturer in Botany, University of Sydney.
(Plate ii; ninety Text-figures. )
[Read 29th March, 1933.

Introduction.

More than thirty years ago Professor D. H. Campbell (1897, pp. 2 and 3)
stressed the fact that the classification of the Angiosperms was little more
than mere guess-work, and in considering a remedy for such a regrettable state
of affairs remarked: “How greatly would the value of many an important mono-
graph on a genus or family be enhanced did it but include a connected account
of the whole life-history of one or two representative forms”; and again, “It is
true that the structure of the flower and fruit of the Angiosperms is of the
greatest importance in their classification, but these alone are not sufficient for
settling positively questions of affinity, except between nearly related groups.
An accurate knowledge of the development and histology of the reproductive
parts and embryo is also very important in this connection. Not until very
much more is known than at present about the life-history of representatives
of all the principal types of flowering plants, shall we be in a position to begin
to build up a system of classification which we can hope to be even approximately
accurate.”

It is significant to observe that almost twenty years later—in 1916—Professor
Pearson (1929, p. 179) in making a morphological comparison of the embryo-sac
and endosperm of Gnetum with those of Angiosperms bewailed the fact that
“Even now, the number of Angiosperm sacs investigated is comparatively small.”
Again, recently Rendle (1925, p. 424) makes the following observation: “The
consideration of the sympetalous Orders together after the dialypetalous is largely
a matter of convenience and must not be regarded as implying that they form a
distinct natural class.”

Complaint then regarding the paucity of information on the details of life-
histories of types representative of the whole Angiosperm phylum is even at the
present day widespread, and the call for further research is clearly insistent.

While some cry out upon the monotonous regularity of the development
of the reproductive parts in Angiosperms, others again are seized with the
idea that therein lies the most valuable approach to the solution of certain
problems of outstanding phylogenetic and systematic importance. Adherents of
the latter view then, need never look beyond such a field for problems worthy
of the most zealous investigation.

The writer, convinced of the real value of any additions to our knowledge
of the several phases of the Angiosperm life-history, has endeavoured during the
last few years to throw some further light on this subject by investigating forms
which are almost confined to Australia (Brough, 1923; 1924; 1927).

Newman (1928, 1929) has also made valuable contributions along similar
lines, and it may be suggested that, with further additions of a like nature,
means may be placed at our disposal, which will permit of the elucidation of
such major problems as the Origin of the Australian Flora, the spread of the

F

34 LIFE-HISTORY OF GREVILLEA ROBUSTA,

Angiosperms in general, and the real relationships existing between the Families
of flowering plants,

This present publication embodies the results of studies in the Proteaceae,
a family which has received much attention—particularly in Australia—from
many investigators specially interested in xerophytes, and their xeromorphic and
edaphic adaptations. This family has been markedly neglected by those engaged
in the investigation of Angiosperm life-histories (except by Ballantine, 1909).

Bews (March, 1927, pp. 6 and 7) furnishes an account of the past and present
distribution of the Family, and draws attention to the diverse views held by
such eminent palaeobotanists as Scott and E. W. Berry regarding its fossil
history and the interpretation thereof.

The Proteaceae comprise 55 genera and some one thousand species (Bews,
May, 1927, p. 69), and a study of the more outstanding systems of classification
shows how incompatible are the views held by past and present-day systematists.
In this connection Bews observes: “According to Engler’s arrangement the
Proteales connect with the Santalales: Hutchinson thinks they are allied with
the Thymeliaceae, Balfour was of the opinion that they had affinities with the
Roseales.” Further, Rendle (1925, p. 58), dealing with the Santalales and
Proteales, remarks: “While undoubtedly allied to each other, it is difficult to
associate these Orders with other groups.”

The obvious deduction from such a number of conflicting opinions is that
the present data are insufficient to settle the questions under dispute. Accordingly
the need for the exhaustive study of the life-history of at least one Australian
representative of the Family need no longer be stressed.

THE PROTEACEAE AND THE PROBLEM.

The Proteaceae form a very conspicuous component of, and in many regions
impart a distinctive facies to, the Australian flora, which embraces some twenty-
nine of the fifty genera and about 575 of the one thousand species, included in
this large Family (Bentham, 1870). Sargent (1930, pp. 577 and 578) points
out that the Australian representatives are massed chiefly in Western Australia,
“Seeing that it possesses nearly twice aS many species as all the other States of
the Commonwealth together’’. He also supplies the following interesting comment:
“Tt seems, therefore, quite beyond dispute that this Family, whose members are
generally regarded as typical xerophytes—drought resisting plants, often, indeed,
drought loving plants—as a Family has a strong preference for the moister
regions of our Island Continent.”

Accordingly it is not surprising to find the Proteaceae well represented in
New South Wales, where some 17 genera and 135 species are located (Moore and
Betche, 1893) amid an environment which cannot be regarded as xerophytic
(Osborn, 1932).

Abundant material for an investigation of the nature proposed is therefore
easily available, and an early consideration was to decide which genus and
species most readily lent itself to the desired objectives. In making a final
selection it was very necessary to keep in mind that the research must primarily
concern itself with an endeavour to glean information regarding (a) the nature
of the perianth, ()) the significance of the nectar scales common—though in
various degrees of size—to members of the Family, (c) the frequent zygomorphy,
(d) the morphology of the gynoecium, (e) the origin, development, and structure
of the ovule, (f) the gametophytes, (g) pollination, (h) fertilization, (i)
embryology, (j) the cause of the pronounced low seed production, and (k) the

i)
a

BY P. BROUGH.

morphological nature of the seed-wing. Other features might be expected to
arise and to demand elucidation as the work progressed.

It seems reasonable to state that detailed information regarding the points
enumerated must decide, or at least very materially aid in determining, the
vexed question of the affinities and systematic position of the Family.

GREVILLEA ROBUSTA (General).

Grevillea, the largest of the Protead genera, comprises 160 species, and of
these 45 are native in New South Wales.

Only as the result of much preliminary investigation was it decided to make
Grevillea robusta—the Silky Oak—the material for intensive study. Many other
types of this and other genera had been considered, but were in turn rejected,
either on account of the early thickening of the ovary wall, or the low percentage
of fertilized ovules or both.

Reference to Plate ii will readily indicate the general features of the form
chosen, which has the advantage of being readily obtainable, has a highly
specialized floral structure, and above all produces a comparatively high per-
centage of seeds with well developed embryos. This last character assumes
enormous importance, when one realizes the notoriously low percentage of seeds
set by the very great majority of the various species of the Family, and the
consequent great difficulty in tracing the embryology. Grevillea robusta is one
of the dendroid representatives of the Proteaceae, and may attain to a height of
120 feet, with a stem diameter of about 8 feet (Francis, 1929, p. 83).

Its natural habitat is in the brush forests, where it ranges from the
Clarence River in New South Wales to Northern Queensland (Maiden, 1917, p.
175). Being a brush type, its range inland is strictly limited, no specimen
having been reported beyond a distance of one hundred miles from the coast.

The Silky Oak, on account of its great ornamental value, has been planted
freely in and around Sydney, so that abundant material is to hand. In sheltered
aspects of this—the Sydney—district anthesis commences about the beginning
of October, and continues until mid-December in the more exposed localities.
In full bloom the tree compels attention, not only on account of the brilliant
orange-yellow of the densely massed inflorescences, but also because of the
abnormal nectar production, which attracts great numbers of bees and numerous
nectar-seeking birds, the ensemble being an extremely lively one from the
naturalist’s point of view.

The inflorescence is a panicle, the individual racemes being three to six
inches long, and varying in position from almost horizontal to nearly vertical.
The raceme supports numerous flowers, each on a pedicel about half an inch in
length (vide photo, Plate ii).

Reference to Text-figure I, 1-27, shows various phases in floral development—
the young bud, the looping of the style, its gradual emergence between two
perianth segments, the final freeing of the stigmatic region, the recurving,
contraction, and fall of the perianth segments, and finally several stages in
development of the fruit. It is to be noted that the style bursts through the
Berianth on the side remote from the axis of the inflorescence, and the tip curves
over adaxially. The perianth segments, bearing the dehisced anthers, as a rule,
fall within a day or two of anthesis, and subsequently a small ovary mounted on
a gynophore, which in turn is supported by the long pedicel, stands exposed.
Anther dehiscence extends over the period just prior to, and simultaneous with,
the escape of the style, and so pollen is available for distribution from the top

36 LIFE-HISTORY OF GREVILLEA -ROBUSTA,

of the expanded style, which bears aloft almost all of the pollen in a cone-shaped
mass. The stigmatic surface is not yet exposed, however (Text-fig. I, 10-11).

It is noteworthy that in the early bud condition the angle between the
pedicel of the flower, and the peduncle of the raceme is acute, but as anthesis
approaches, this angle increases to about 90° and later may become very oblique.

Text-fig. I.
Series 1-27 illustrates the development of the flower-bud, flower, fruit and winged
seed of Grevillea robusta; the follicular nature of the fruit, and its method of dehiscence

are also depicted. x 1.
Series a-f shows successive stages in the production of a fruit from a “capped”

flower of Grevillea Banksii. x 2.
Series A and B illustrates fruit production from ‘‘capped’”’ flowers of Grevillea

robusta. x 3.

BY P. BROUGH. 37

This change obviously places the flowers in more advantageous position for
pollination, since birds alight on the stronger basal region of the inflorescence—
the distal end being too fragile for their support. The open flowers thus
conveniently face the pollinating visitor.

Floral Organogeny.

In order to study the origin and early development of the floral members,
transverse and longitudinal microtome serial sections of complete young inflores-
cences and also of mature individual flowers were examined.

A raceme in longitudinal section, as illustrated in Text-figure 1, consists of an
elongated central axis or peduncle on which are arranged, in acropetal order, the
numerous floral buds, which, as is clearly demonstrated, arise in pairs and all
round the axis (Text-fig. 2), each pair being located in the axil of a single hairy
bract (Text-figs. 1, 2, 3 and 5) which is early deciduous. This twinning of the
flowers is readily perceived in the genus Banksia, even in mature inflorescences,
but in Grevillea robusta this feature tends later to be obscured owing, in the first
place, to the heliotropic twisting of the pedicels and, secondly, to the fall of
many young buds. In this form no exception to the paired arrangement of the
flower buds was encountered, though numerous serial sections were examined.

The first evidence of the flower is a rounded mass of tissue (Text-fig. 1 at A)
on which, as development proceeds, four symmetrically arranged protuberances
are recognized. Two of these are evident in Text-figure 1 at B, and such represent
_the primordia of the perianth segments. A bract subtends each pair of buds
(Text-fig. 2).

By following the peduncle basipetally in Text-figure 1 the gradual increase in
size of the perianth segments may be traced, and the inception of a second and
inner set of four primordia recognized. The illustration in Text-figure 4 shows
such primordia at a more advanced stage, and their further development is
depicted in Text-figures 4-9 inclusive.

The primordia referred to, which are situated directly in front of and
within the perianth segments, are those of the four stamens, each of which
becomes epiphyllous at a very early stage of development, because very soon
after their inception, the meristematic region of each perianth segment fuses
with that of the stamen immediately within (Text-fig. 4), so that the two fused
fioral members continue growth as a single structure until the mature condition
is attained when the anther, supported on, and in conjunction with, a very short
part of the filament, is the only free part of the stamen (Text-fig. 9). |

It will be noticed (Text-fig. 1, at B) that a vascular strand passes from the
stele of the peduncle to each of the subtending bracts. Again, vascular tissue
runs to the base of each pair of flower buds, at the base of which it bifurcates,
one fork passing to each flower. Hach of these bundles then divides into five,
one branch passing into each perianth segment and the base of the organic apex
respectively (Text-fig. 5), and each of these bundles in turn splits into two (Text-
fig. 1, at C) one portion supplying the conjoined stamen while the other proceeds
towards the distal end of the perianth segment.

Again, reference to Text-figures 4-9 shows that the primordium of the single
carpel makes an early appearance. Its true nature is, however, best illustrated
in a series of transverse sections of buds at successively more advanced stages in
development (Text-figs. 12-14), where the fact that the gynoecium consists of a
single carpel—clearly an infolded simple _ leaf-like structure—is amply
demonstrated.

38 LIFE-HISTORY OF GREVILLEA ROBUSTA,

J
SS i
SS

i
it
Hf
|
i!
,

Text-fig. 1*—A median longitudinal section of a very young inflorescence
illustrating various phases in development of the flower bud and subtending
bract. The vascular system of the peduncle, v.s., and its continuity with that
of the buds and bracts is indicated. x 35.

BY P. BROUGH. 39

At the very young stage depicted in Text-figure 12, a midrib with two lateral
branches is evident, and at a slightly older stage two additional bundles make
their appearance (Text-fig. 13). Further increase in size of the carpel results in
a marked increase in the number of vascular tracts until, at the stage represented
in Text-figure 14, as many as 13 bundles may be observed. A marginal bundle
passes into the funiculus of each of the young ovules depicted in Text-figures
14 and 26. Also, in Text-figure 12, the incipient marginal placentae which later
give rise to the two ovules of the carpel are just evident at X, while in Text-
figures 15, 16, 26, 33, 34, 57 and 58, the origin and general structure of the young
ovules are shown.

Again, very soon after the gynoecium is initiated, an upgrowth of paren-
chymatous cells from the torus is noticeable at the base of the carpel and on
the adaxial side. This tissue gradually increases in height, and later forms a
conspicuous crescent-shaped nectar scale. It is well developed even in such an
early bud stage as is illustrated in Text-figures 7-9.

All the floral organs have thus become established, and it only remains to
follow their development until maturity of the flower is attained. The position
of each member relative to the axis of the inflorescence will also require careful
elucidation.

In order to gain a proper appreciation of what may be regarded as the
essential floral structure of Grevillea robusta, a series of transverse sections (Text-
figs. 17-32) and also longitudinal sections (Text-figs. 33 and 34) at the stage just
prior to anthesis were prepared. An appreciation of the general structure of the
flower bud, as shown in Text-figure 33, will readily indicate the particular levels
at which the various transverse sections of the series were made.

Examination of selected transverse sections taken at intervals and in acropetal
order indicates a shallow concavity in the floral receptacle. Thus the flower is
slightly perigynous. Additional evidence of this is supplied by a study of longi-
tudinal sections of buds at various stages of development (Text-figs. 9, 11, 33 and

Text-fig. 2.—A transverse section of an inflorescence cut about the region
C in Text-fig. 1. The twinning of the floral buds with subtending bract, b., and
the vascular system are shown. x 35.

Text-fig. 3A median longitudinal section of two slightly older buds
showing perianth segments, p. x 35.

Text-fig. 4-—A median longitudinal section of a still older flower depicting
the young anthers, st., and the central upgrowth of carpellary tissue, C. x 35.

Text-figs. 5, 6.—Longitudinal sections of twin flower buds slightly more
advanced than in Text-fig. 4. A vascular supply from the peduncle to each
bud, and a faint lobing of the carpel are shown; b., bract; p., perianth segment ;
St., stamen. x 35.

Text-fig. 7—Median longitudinal section of flower-bud cut at right angles
to the antero-posterior plane. Microspore mother-cells are present in the anther,
st.; two young megasporangia, m., are seen in the ovarian cavity, o.v., and the
nectar-scale, n.s., is visible at base of carpel. x 35.

Text-fig. 8.—Median longitudinal section of flower-bud cut in antero-posterior
plane. n., nucellus; f., filament of anther; m.m., microspore mother-cells;
n.S., nectar-scale; i.p., interlocking perianth segments; s., style. x 35.

Text-fig. 9.—Longitudinal section as in previous figure, but at a slightly
older stage: The microspore mother-cells are undergoing reduction division. x 35.

Text-fig. 10.—Transverse section through apical region of flower-bud:
p., perianth segment with epiphyllous stamen, st., containing four loculi; az.,
axis of inflorescence; s., style. x 50.

* All drawings, with the exception of that in Text-figure 83, are camera
lucida.

LIFE-HISTORY OF GREVILLEA ROBUSTA,

40

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BY P. BROUGH. 41

34). From the torus the perianth segments, stamens, nectar-scale and carpel
gradually differentiate centripetally. At first the perianth segments are in close
contact, but the subsequent extraordinary growth in length of the style neces-
sitates its curvature, which takes place in the antero-posterior plane. With
further increase in its length a decided loop is formed which forces all the
segments slightly apart, but the gap between the two abaxially placed perianth
segments is steadily emphasized, and it is through this aperture that the loop
of the style eventually is thrust (Text-fig. 31; also Text-fig. I, 6-9).

The series of transverse sections depicted in Text-figures 17 to 32 illustrates
the structure of the flower at various levels. The lobed carpel, the adaxial nectar-
scale, the relative position of the separate perianth segments, the ovarian cavity,
the marginal position of the young ovules, the style—in part within and in part
without the perianth—the fusion of the stamens with the perianth segments, and
the cone-shaped nature of the apex of the style are all clearly delineated.

In Text-figure 32 of the series the apical part of the flower is seen in longi-
tudinal section, this of course being due to the curved nature of this region, which
is approximately at right angles to the axis of the flower as a whole. At this
stage the anther contains tetrahedral pollen grains.

The carpel is so orientated that the line of fusion of the margins is adaxial
and adjacent to the central region of the nectar-scale (Text-fig. 22). The position
of these organs relative to the perianth segments is also demonstrated.

It is to be noted—despite statements to the contrary—that the primordia
of the perianth segments are separate from the first, and that the fused condition
does not exist at any stage of their development. None the less it is evident that
in the young bud the margins of adjoining perianth segments fit into each other
with extraordinary closeness and precision by means of interlocking teeth-like
cells (Text-figs. 8, 10 and 35). The rigidity of this structure results in a simula-
tion of the sympetalous condition.

In the young bud condition then, the style keeps pace with the development
of the perianth segments, but subsequent restriction in rate of growth of the
perianth does not mean a similar slowing down in style elongation, which
steadily proceeds. It might be expected that the style should force its way out

Text-fig. 11.—Median longitudinal section of lower part of a flower-bud cut
as in Text-fig. 7, showing carpel, ovarian cavity, two young megasporangia,
and nectar-scale. x 225.

Text-fig. 12—A transverse section of a young flower-bud showing inner
margins of perianth segments, p., enclosing a very young carpel with infolded
margins; v.b., vascular bundles; «#, meristematic region giving rise to placenta
and nucellus. x 135.

Text-fig. 13.—A transverse section of a bud slightly older than in previous
figure. The epidermal cells of the infolded margins of the carpel and an increase
in the number of its vascular bundles are depicted. m., megasporangium ;
v.b., vascular bundle. x 135.

Text-fig. 14.—A transverse section of a still older bud showing the vascular
bundles of perianth and carpel respectively. The two megasporangia, m. and
m,, seated on the margin of the carpel are evident. A vascular bundle is seen
to pass from the margin of the carpel into the ‘chalazal region of each nucellus.
XD:

Text-fig. 15.—Vertical section in lateral plane through the ovarian cavity
containing two young megasporangia. xX 225.

Text-fig. 16.—Median vertical section in antero-posterior plane through a
young nucellus. The origin of the incipient inner integument is indicated at X.
The embedded megaspore mother-cell, m.c., is evident. x 225.

LIFE-HISTORY OF GREVILLEA ROBUSTA,

Text-fig. 17-32.—A selection of transverse sections taken from a complete
acropetal series cut through a flower bud at the stage of development indicated
in Text-fig. I, 5. These sections illustrate the general structure and relative
positions of the component parts of the flower. Sections 17-21 are slightly
oblique to axis of flower. For detailed description consult text. c., carpel;
p., perianth segment; wn.s., mnectar-scale; yn.s.l., mectar-scale lobes; ™m.,
megasporangia; f., funiculus; s., style; st., stamen; s.d., slit of dehiscence;
m., microspores; Aw., axis of raceme. x 25.

BY P. BROUGH. 43

YY)

St DD Ax

32

through the perianth segments at the apex of the flower, but in this region, the
interlocking of the cells, which is specially marked at the distal ends of the
perianth segments, exerts sufficient resistance to overcome the upward thrust,
with the result that the middle region of the style curves and eventually forces
its way out between the two abaxial segments, where there must be less
resistance, even though interlocking occurs in this region also. Examination of
the style shows elongation of the cells on the outside of the curve, and compres-
sion of those on the convex face. However, this steady increase in length of
the style imposes a very considerable and ever increasing strain, with the result
that the distal end of the style eventually frees itself by forcing its way through
the interlocked ends of the perianth segments, which, thus relieved suddenly
from an elongating tension, contract. They then appear as somewhat incon-

44 LIFE-HISTORY OF GREVILLEA ROBUSTA,

spicuous coiled structures at the base of the floral axis (Text-fig. I, 10-11). There-
after they are soon deciduous—only persisting for a few days.

Investigation of sections cut longitudinally through the flower bud (Text-

fig. 8) shows a well-defined absciss layer of cells (Text-fig. 36) just adjoining the
base of the perianth.

The line of fracture is visible in Text-figure I, 7-10.

Thus, perianth fall in
regard to time and tissue concerned is determined with precision beforehand.

Now, although the above description represents the normal behaviour of the
flower at this stage of development, it is interesting and significant to record that
in some cases the apex of the style does not forcibly free itself by a sundering

D

fa
ie,

Le

ft

33

Text-fig. 33.—Median longitudinal section of flower bud at stage
in Text-fig. I. f.r., floral receptacle;

2 of series

n.S., nectar-scale ;
c., nectar-secreting cushion between nectar scale and perianth; o.v., ovule;
p., perianth; s., style; v.b., vascular bundle; s.t., stigmatic tissue; a., anther.
x 25.

a.l., absciss layer;

Text-fig. 34.—Median longitudinal section of lower part of a bud slightly

older than, and cut at right angles to, that shown in previous figure. Inter-
pretation as in Text-fig. 33. x 25.

BY P. BROUGH. 45

of the interlocked distal ends of the perianth segments, but instead, such segments
are ruptured at their individual bases, along the absciss layer, and are carried
up on top of the style (Text-fig. I, A, B) just as occurs in such well-known cases
as the corolla in Vitis and the staminate flowers of Casuarina. It therefore follows
that in these particular cases the resistance exerted by the interlocking at the
apex of the perianth segments is great enough to force a prior rupture at the
bases of these segments. When this happens, the force tending to sunder the
apices of the segments is destroyed, with the result that the perianth becomes
lodged as a “cap” on top of the style until long after the pollination period
has elapsed. Further discussion regarding this phenomenal behaviour will be
deferred until the general question of pollination is dealt with.

In the normal flower anthesis is completed by the sudden disengaging of the
distal region of the style from the enclosing perianth segments.

The monocarpellary ovary has a bulbous-shaped hollow ovary supported on a
gynophore, and surmounted by a long curved style (Text-fig. I, 12-25). The apex
resembles a low cone with a blunt top which, at the moment of liberation, is piled
high with pollen. Immediately subsequent to this stage, the cells occupying the
central region of the distal end of the style grow rapidly, rupture the surmount-
ing epidermis, and form a loose mass of elongated cells disposed around a funnel-
like opening (Text-figs. 33 and 34). This cavity gradually widens and exposes
the elongated glandular cells of the stigma (Text-figs. 37, 38 and 39). At the
same time this central upgrowth displaces centrifugally any microspores which
have not been dispersed, and causes them to be removed from possible contact
with the stigmatic cells.

The flower thus shows extreme protandry, and enforces cross pollination. The
mature stigmatic tissue, when examined in longitudinal and transverse section
respectively, is seen to be composed of long vacuolate cells (Text-fig. 39), which,
lining a deep pocket (Text-fig. 37), furnish an ideal medium for the development
of the male gametophyte.

Nectar Secretion.

Profuse nectar secretion is a characteristic of the Proteaceae, the flowers of
which are provided with nectar-secreting organs of very varying form. When
Jarge and separate from one another, as, for example, in the genus Banksia,
they are seen to alternate in position with the perianth segments.

In Grevillea robusta, however, the scale is a single crescent-shaped structure
situated at the base of, and forming a partial collar around about two-thirds of the
circumference of, the gynophore (Text-fig. 22). Owing to two shallow clefts in
its central region, the scale is provided with two lateral broader lobes and a
narrower central lobe (Text-fig. 24). It measures 0-5 mm. at its greatest height,
is located on the adaxial side of the gynophore, and gradually tapers away on
approaching the abaxial side. The scale is markedly thickened at the base on
the adaxial side, where an additional but much smaller cushion of tissue is
located (Text-figs. 40 and 43). Microscopic examination of the adaxial and abaxial
surfaces of this scale and cushion shows that the greater part of the area is com-
posed of protruding secretory cells, either isolated or in groups of three or four,
imparting to the whole surface a modified botryoidal appearance (Text-figs. 40, 41
and 43). These cells stain very deeply with safranin or Haidenhain’s iron-alum
haematoxylin stains. Clearly then this scale, with its adjoining cushion, is a
nectar-secreting organ.

46 LIFE-HISTORY OF GREVILILEA ROBUSTA,

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fEASE ite :

AL SNe ’
SEH?
eee Hee
@iern y def

jieseess eel am
SESE
ST SH
KD CALLAN
EE We OS.
Pane gerenee tas UA of
SEER
Saas 9,
ont
ai

SNS REE.

BY P. BROUGH. 47

But secretion is not confined to this region, for an investigation of the basal
part of the perianth segments in longitudinal section (Text-figs. 40 and 42) and
in surface view shows that the majority of the epidermal cells protrude as finger-
like processes, their length in many cases being several times their breadth.

Such cells do not take up a position at right angles to the perianth segment,
but are applied for the most part to its surface in an irregular fashion. This
perianth secretory region extends from the base of each segment to a height of 1:5
mm., so, taking into account the great volume of the protruding cells as well
as the extensive area bearing them, it seems certain that the amount of nectar
derived from this region must considerably exceed that which has its origin in
the scale.

The combined areas of the secretory cells referred to are relatively very
considerable, and their activity readily accounts for the exceptionally large pro-
duction of nectar in the individual flower. Thus the nectar-scale is not—contrary
to expectation—the sole or even the main source of nectar secretion, but rather
plays a subsidiary role in this regard.

On the adaxial side of the flower a natural reservoir for holding the nectar
is formed between the gynophore and the adhering bases of the opposing perianth
segments, and also between the gynophore and the scale (Text-figs. 23 and 40).

Morphology of the Nectar-Scale.

The fact that the nectar-scales are, in this Family, often four in number,
and alternating with the perianth segments, suggested to the writer that they
might constitute a reduced and modified inner whorl of floral leaves, or in other
words represent the vestiges of an ancestral normal corolla, in which case the
present perianth would fall to be interpreted as the calyx.

In an endeavour to settle this question, which turns on the morphology of the
nectar-scales, series of transverse (Text-figs. 22-24) and longitudinal sections
(Text-figs. 33, 34, 40 and 43) of flower buds at successively older stages of
development were examined. In every case the scale and cushion were found to

Text-fig. 35.—Transverse section through portions of two adjoining perianth
segments, shoying interlocking marginal cells, i.c., and nectar-secreting cells,
N.C. X 225.

Text-fig. 36.—A longitudinal section showing cells of absciss layer, a.l., at
base of perianth segment, p. x 225.

Text-fig. 37.—Median longitudinal section of the cone-shaped apex of the
style. The papillate nature of the stigmatic cells lining the central funnel-
shaped cavity is depicted. Some of the receptive stigmatic cells are protruding
beyond the rim of the cavity. x 135.

Text-fig. 38.—Transverse section of stigmatic cavity. x 225.

Text-fig. 39.—Typical form of a stigmatic cell; the vacuoles are apparent.
xDD0s

Text-fig. 40.—A longitudinal section through a nectar scale, a2.s., and base
of a perianth segment, p. The protruding nature of the secretory cells covering
the whole of the former, and lining the inner surface of the latter are indicated;
Ss, style. x 38.

Text-fig. 41.—Longitudinal section of the marginal region of a nectar-scale
showing secretory cells, s.c., of epidermis. x 225.

Text-fig. 42.—Longitudinal section cut in the region of the ovary showing
nectar-secreting epidermal cells, s.c., of perianth adjoining epidermis, e., of style.
x 225.

Text-fig. 43.—Longitudinal section of cushion of tissue at base of nectar-
scale showing general nature of the nectar-secreting cells, s.c.; a few of the
cells of the absciss layer, a.l., are evident. x 225.

48 LIFE-HISTORY OF GREVILLEA ROBUSTA,

LO

Ol

2
pena
feacSt

CT
LPH

May a
1 wel

Text-fig. 44.—Transverse section of young flower bud showing portion of
two perianth segments, p., two adjacent anthers in part and the central style, s.
sp.c., Sporogenous cells; w., wall cells; v.b., vascular bundle. x 135.

Text-fig. 45.—Transverse section of young anther slightly older than in
preceding figure. ep., epidermis; ¢., tapetum; w., intermediate wall layers;
S.m., microspore mother-cells. x 225.

BY P. BROUGH. 49

consist entirely of thin-walled parenchymatous cells without any differentiation,
except for the papillate nature of the epidermal secretory cells already discussed.
There is no indication, whatsoever, of any associated vascular tissue, and con-
sequently no ground for interpreting the scale as the morphological equivalent of
a reduced perianth. It would seem then to be merely an adventitious accessory
development of the torus.

The Microsporangium: Development and Structure.

The anther consists of four microsporangia held together by the connective
tissue of the filament. Examination of sections of anthers at various stages
showed that in the main the development conforms to what may be termed normal.
Nevertheless, features of peculiar interest are encountered. For example, the
archesporium becomes differentiated in the microsporangium of a flower bud about
the stage represented in Text-figure 4. It is diagnosed as a plate of hypodermal
cells, one cell thick, extending practically throughout the whole length of the
anther. These cells are slightly radially elongated, possess large nuclei and
have dense cytoplasm. Later, each of the archesporial cells divides by a peri-
clinal wall, thereby giving rise to a primary parietal and a primary sporogenous
layer respectively. The former, by active periclinal and anticlinal divisions,
initiates a wall four cells thick lined internally by a fifth layer, the tapetum
(Text-figs. 44, 45), easily recognized by its large, radially elongated, and
densely staining cells. Meantime the primary sporogenous layer remains almost
quiescent, the only notable change being an increase in the size of the constituent
cells which, however, do not normally divide in order to produce an increased
number of sporogenous cells, but, instead, function directly as spore mother-cells.
It may be mentioned that occasionally an individual primary sporogenous cell
does divide to give rise to two derived sporogenous cells, but such is a rare
oecurrence.

At first the spore mother-cells are closely packed and polygonal (Text-fig. 45),
but, with the enlargement of the sporangial cavity consequent on a general increase
in size of each microsporangium, they become free from one another, assume a
rounded form, and float in the nutrient medium produced by the enlarged,
actively secreting and now multinucleate cells of the tapetum (Text-fig. 46).

A typical microspore mother-cell is figured in Text-figure 49, where the
characteristic large nucleus, coiled chromatin, nucleoli, dense cytoplasm and thin
wall are evident.

Later the condition known as synizesis (Text-fig. 50) was diagnosed, and in
slightly older material the various phases in reduction division were traced. The

Text-fig. 46.—Longitudinal section of a microsporangium at a still older
stage. ep., epidermis; ¢., tapetum; w., wall layers; s.m., free microspore mother-
cells. x 225.

Text-fig. 47.—Tranverse section of two adjacent anthers at the stage just
prior to dehiscence. The tissue separating adjoining microsporangia has been
ruptured, and the region of the slit of dehiscence is evident. Portion of two
perianth segments is shown; m., tetrahedral microspores; v.b., vascular bundle of
connective. x 55.

Text-fig. 48.—Transverse section of part of each of two adjacent anthers.
The rupture of the tissue separating two microsporangia of an anther is indicated
at w. The epidermal cells have collapsed at this stage, and the fibrous layer,
f.l., now delimits the sporangium. The overlapping unthickened terminal cells,
o.c., of the fibrous layer, which later split apart and thus provide a longitudinal
slit of dehiscence, are delineated. m., tetrahedral microspores in binucleate
condition. x°225.

50

LIFE-HISTORY OF GREVILLEA ROBUSTA,

Text-fig. 49.—Microspore mother-céll as in Text-fig. 46. x 1,000.

Text-fig. 50.—Slightly older microspore mother-cell in stage known as
synizesis. x 1,000.

Text-fig. 51.—Section of microspore mother-cell cut at right angles to the
reduction division spindle in the late anaphase. The individual chromosomes,
10 in number, are shown. x 1,000.

Text-fig. 52.—Microspore mother-cell containing spindles of two nuclei
dividing simultaneously. -x 1,000.

Text-fig. 53.—Three microspores of a tetrad still contained within wall
of mother-cell. x 1,000.

Text-fig. 54.—Two adjacent pollen grains in sitw in a microsporangium.
The walls have become much thickened. The spore on the left has reached
the binucleate condition, while the nucleus of the other is in the metaphase of
division. In section the shape of the spore depends on the plane in which it
is cut. x 1,000.

Text-fig. 55.—A pollen grain at the binucleate stage. Only one nucleus
containing a large nucleolus is present in this section. The other body, which
simulates a nucleus, is a vacuole containing deeply staining bodies. x 1,000.

Text-fig. 56.—Mature binucleate microspore at dehiscence stage of pollen
sac. <A thick exine with outer layer of cutin, and the thin intine, protruding at
the corners of the spore, are indicated. At this stage a thin membrane separates
the smaller generative cell, g., from the larger tube nucleus. x 1,000.

BY P. BROUGH. 51

number of chromosomes in the haploid condition was determined as ten, this
number being specially well demonstrated in sections cut at right angles to the
spindle in the late anaphase (Text-fig. 51). At this period the wall of the spore
has increased in thickness. The daughter nuclei divide immediately after forma-
tion, the spindles of this phase—one of which is oblique—being shown in Text-
figure 52. Each of the resulting nuclei then becomes enveloped in a quarter-share
of the cytoplasm, which in turn is bounded by a thin membrane, and thus the
tetrad condition is attained (Text-fig. 53).

While microsporogenesis is proceeding, certain well-marked changes fall to be
recorded in the development of the tapetum and wall of the sporangium. ‘To
begin with, the cells of the hypodermal layer become characterized by an
increasing number of starch grains, which later furnish material for the pro-
nounced fibrous thickenings characteristic of this layer at a more mature stage
(Text-fig. 48). Meanwhile the tapetal cells enlarge (Text-fig. 46), become multi-
nucleate, and gradually absorb the wall layers within the endothecium. However,
after the inception of the microspores, the vitality of the tapetum steadily
decreases, and by the time anther dehiscence is imminent, only a few frag-
mentary remains mark a hitherto prominent feature of the sporangium.

Another change, contemporaneous with the decline of the tapetum, is the
collapse and gradual disappearance of the somewhat thin-walled and fragile cells
of the original epidermis, so that, in the mature microsporangium (Text-figs.
47, 48), the fibrous layer delimits, and, for all practical purposes, constitutes the
free portion or arms of the sporangial wall.

A similar occurrence has been recorded in Dampiera stricta (Brough, 1927,
p. 480), while in Styphelia longifolia (Brough, 1924, pp. 166 and 167) the fibrous
layer is morphologically the epidermis. This progressive relegation of the endo-
thecium to the outside of the mature sporangium is no doubt related to the hot,
dry conditions commonly prevailing at the time of anthesis in many Australian
fiowers. Reference to Text-figure 47 shows the general structure of the anther
just prior to anthesis.

The partition wall separating the pairs of microsporangia has ruptured,
resulting in the formation of two pollen sacs in each anther. An examination of
the fibrous layer indicates a somewhat noteworthy and peculiar dehiscence
mechanism. It is observed that at o.c. in Text-figure 48, where the endothecial
layer of one microsporangium meets the corresponding layer of the adjoining
sporangium, a double layer of cells devoid of fibres is formed by the overlapping
of the tapering ends of the respective endothecial layers. As usual, then, a region
of weakness extends throughout the length of the pollen sac, but at dehiscence,
instead of the rough fracture, so commonly found in anthers, there is an orderly
separation of the cells along the line of the overlap, and a subsequent pulling
apart of the two freed arms, each terminated by a row of unbroken tapering
cells—a somewhat unique phenomenon in anther dehiscence (Text-fig. 32).

The Male Gametophyte.

After the uninucleate microspores have been set free by the dissolution
of the wall of the mother-cell, they rapidly increase in size and gradually assume
a precise tetrahedral form in the nearly mature flower bud (Text-figs. 48, 56). This
shape is characteristic of most Protead forms. Soon after this, but before anthesis,
the nucleus undergoes mitotic division, and the binucleate stage is attained, the
two nuclei being separated by a thin membranous wall (Text-figs. 54, 56). The

52 LIFE-HISTORY OF GREVILLEA ROBUSTA,

generative nucleus is the smaller and is more densely encased in cytoplasm than
the larger centrally placed vegetative nucleus.

Meanwhile, the wall becomes excessively thickened, the massive exine being
differentiated into two layers, a clear outer covering of cutin and a thicker
and deeply staining inner layer. A thin membrane represents the intine (Text-
fig. 56). This thickening of the exine is not uniform, however, and is distributed
in such a manner that four unthickened areas are evident at the points of the
tetrahedron.

Just prior to anthesis one or more vacuoles make their appearance in the
cytoplasm of the pollen grain. In preparations stained by Haidenhain’s iron-
alum haematoxylin method it was observed that such vacuoles contain numerous
deeply-stained granules (Text-fig. 55) which may be interpreted as reserve food
material—a useful adjunct in the nutrition of the relatively long pollen tube. As
such vacuoles are of large size relative to the microspore as a whole, the con-
sequent increase in internal pressure is no doubt the cause of the protrusion
of the intine at the four corners of the tetrahedron (Text-fig. 56). Such vacuolation
seems to be a constant prelude to spore germination, for it is at this stage of
development that the style frees itself from the perianth, and raises the pollen
aloft on its distal extremity.

On germination, one of the protuberances grows out to form a pollen tube,
and its subsequent development agrees closely with that previously described in
the case of Dampiera stricta (Brough, 1927, p. 482). It may be added that the
thickening of the spore wall is more pronounced than in Dam~piera.

Examination of the receptive stigma in longitudinal section revealed pollen
grains in situ. These germinate, as indicated, and subsequently pollen tubes
traverse the long style and eventually were observed to enter the ovarian cavity,
at the chalazal region of the two pendulous ovules (Text-fig. 34). The pollen
tube pursues its course downwards, adhering to the thin-walled slightly papillate
cells lining the ovary wall, and evidently deriving nourishment therefrom. The
space between the wall and the ovules is just sufficient to permit easy passage
of the pollen tube which, on reaching the lower end of the ovarian cavity, turns
almost at right angles and thereby comes into contact with the apical region
of the pendulous ovule, along which it grows until the neighbourhood of the
micropyle is reached. The micropyle is a relatively wide funnel-shaped passage,
easy of access. The cells of the inner integument bordering the upper region of
the micropyle are of a strikingly papillate nature and admirably adapted to stimu-
late further growth of the pollen tube, which eventually penetrates the crushed
cells of the nucellus, and enters the embryo sac in the normal way.

The Ovule.

The gynoecium is represented by a single carpel occupying the organic apex
of the thalamus (Text-figs. 5-9). Transverse and longitudinal sections both
demonstrate the single foliar nature of this organ which is therefore mono-
carpellary.

The vascular supply in the very young carpel is at first represented by a
single median vascular strand, but as development proceeds lateral veins arise,
which in turn branch, and eventually send vascular extensions to the carpellary
leaf margin (Text-figs. 12-14).

At a very young stage a slight protuberance is noticeable near the base of
each of the infolded margins of the carpel. This is due to the meristematic

on
ew

BY P. BROUGH.

activity of the densely cytoplasmic cells of this region. Evidently such expan-
sions represent the combined young placentae and incipient nucelli, the latter of
which protrude slightly into the ovarian cavity (Text-fig. 12, at x). There is
no very decided placenta, a fact which is attested by the study of similar sections
through older material (Text-figs. 13, 16 and 33). In Text-figure 13 the impinging
but distinct epidermal layers of the inrolled margins of the carpel are clearly
demonstrated. Gradually the nucellus and short funiculus become differentiated
(Text-figs. 13, 15 and 33) and a slight curvature (Text-fig. 16), indicative of the
final anatropous condition, becomes manifest. The integuments, however, are still
undeveloped. As growth proceeds the curvature of the ovule becomes more
apparent, and is steadily accentuated, until the relatively mature phase demon-
strated in Text-figures 57 and 58 is arrived at,

5 OO b/-
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Se

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Fok. Ao ae) t
EOE T
ES

ano:

Co}

Sem

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Ss eo eicoued
[coh

Text-fig. 57.—Median longitudinal section cut in antero-posterior plane of
the lower part of a flower bud showing ovule in ovarian cavity. p., perianth;
s., Style; v.b., vascular strand of carpel; v.s., vascular strand from carpel
passing to chalaza of ovule, 0.; 0.c., ovarian cavity; .s., nectar-scale. x 55.

Text-fig. 58.—Median longitudinal section of an ovule showing general
structure and functional megaspore which later develops into the female
gametophyte. o0.c., ovarian cavity; ch., chalaza; v.b., vascular bundle; o0.i.,
outer integument; i.i.. inner integument; f.m., functional megaspore; 7., nucellus;
p.c., papillate cells of nucellus. x 225.

Herein is evident the essential gross structure of the young ovule—the short,
thick funiculus with central vascular bundle, the massive nucellus, and two
partially developed integuments.

Now, the mature flower is bilaterally symmetrical, and the margins of the
carpel fuse on the adaxial side. The two ovules of each ovary are right and left
of the median plane of symmetry which is antero-posterior. Accordingly, in the

54 LIFE-HISTORY OF GREVILLEA ROBUSTA,

case of longitudinal sections the cut must be made at right angles to this plane,
if both ovules are to be exposed in a single section (Text-figs. 11 and 34). How-
ever, in addition to this, there is the fact that the somewhat flattened ovules are
orientated to lie in the antero-posterior plane, so that this plane must be followed
in section when longitudinal median sections of the ovule are required. Such a
section will of course only pass through one ovule at a time (Text-figs. 33, 57, 58).

A noteworthy feature of this particular ovule is the relatively late inception
and slow development of the integuments. For example, at the stage when the
megaspore mother-cell has become differentiated (Text-fig. 16), there is merely
a slight indication at x of the region of origin of the inner integument. Even at
a much older stage (Text-fig. 58) where the functional megaspore has become
apparent, the integuments are far short of completely enclosing the nucellus,
and it is not until the eight-nucleate stage of the female gametophyte has been
realized, that the nucellus becomes completely invested. At the stage of develop-
ment indicated in Text-figure 58, the outer integument is two, and the inner
integument three cells thick, but as maturity of the female gametophyte is
attained, the inner integument typically increases to five cells broad towards its
apex, while the outer integument increases to eight cells in thickness (Text-fig. 71).
So far then, the integuments are comparatively thin structures. ;

The nucellus is of the massive type, and the manner of its development by
the very regular formation of periclinal and anticlinal walls is indicated by the
uniform arrangement of its constituent cells (Text-figs. 16 and 58). The exposed
epidermal cells of the nucellus are distinctly glandular in appearance (Text-fig. 58),
and remain so until after fertilization. This structural feature is, as already
mentioned, also characteristic of those cells of the inner integument which line
the micropyle, especially those bordering the rim.

It is difficult to dogmatize on the particular réle of such cells, but it may
be that they secrete a liquid whose chemotactic influence guides the pollen tube
direct to the micropyle. In any case they are eminently suited to facilitating the
progress of the pollen tube towards the embryo sac.

Ballantine (1909, p. 161) calls attention to the occurrence of such cells in
Protea Lepidocarpon, and their presence in the ovules of different genera, and
from plants peculiar to two such widely separated continents as Africa and
Australia, is both interesting and suggestive.

Subsequently, the ovule increases greatly in size, the increment being
primarily due to the extraordinary activity of a zone of meristematic cells
situated in the chalazal region of the ovule. The ovule steadily increases in all
dimensions, but the most noteworthy feature of this later development is the
marked increase in length of the ovule, which now assumes an elongated,
flattened form.

Megasporogenesis.

Ballantine (1909) has already furnished, inter alia, an account of the develop-
ment of the megaspore mother-cell, spore tetrad, functional megaspore and embryo
sac of a Protead form, Protea Lepidocarpon, and in consequence the present writer
will only present for purposes of comparison the more significant features of
these particular phases of the life-history of Grevillea robusta.

At the stage in development of the nucellus indicated in Text-figures 16 and 26
respectively, the megaspore mother-cell is recognized by its relatively greater size
and large nucleus. It is also depicted in transverse section in Text-figures 59 and 60.
Although only one such cell is fully differentiated, it is to be noted that some

ao
ol

BY P. BROUGH.

of the contiguous cells are slightly larger and more densely cytoplasmic than the
remainder of the cells of the nucellus. The nature of these cells surrounding the
mother-cell suggested that there might be a plurality of mother-cells in Grevillea
robusta, but careful search failed to reveal any evidence of such. Ballantine (1909,
p. 161) refers to this feature when he observes, “a small group of large cells
situated below the hypodermal layer includes one which becomes the megaspore
mother-cell”. The mother-cell is deeply sunken, and occurs in what may be
termed the distal region of the sporogenous tissue, as opposed to the purely
sterile region of the regular wall layers.

The development of the megaspore mother-cell results in the formation of
four megaspores (Text-figs. 61, 62) which at this stage are arranged in descending
order of magnitude, the largest adjoining the chalazal region of the ovule. The
spores are separated by thin walls. The innermost spore steadily increases in
dimensions, and eventually supplants those more superficially placed (Text-fig..63).

The germination of the functional megaspore (Text-figs. 58, 64) is heralded
by a marked increase in volume, while vacuoles become increasingly apparent
in the cytoplasm. The nucleus divides to give the binucleate stage of the female
gametophyte (Text-fig. 65), and thereafter development proceeds along normal
lines until the eight-nucleate stage is attained (Text-fig. 66). Meanwhile the
sac has increased in all dimensions, and continues to do so, but more particularly
in length, until at maturity (Text-figs. 70, 71) it assumes a much elongated
cylindrical form, a shape which is in conformity with the elongating nucellus.

The Female Gametophyte.

The mature gametophyte is depicted in Text-figure 66. The egg-apparatus
consists of three cells arranged in typical fashion. Each of the two synergids
is pyriform in shape, has a large central vacuole, and contains a relatively small
nucleus. The ovum about the time of fertilization is encased in an aggregation
of starch grains (Text-fig. 68). The polar nuclei first come into contact slightly
to the micropylar side of the middle of the embryo sac, but later change their
position, actual fusion—various stages of which were observed—occurring in
the immediate neighbourhood of the ovum (Text-fig. 69). The antipodal cells are
three in number, separated by walls and located as usual in the lower and some-
what narrowed end of the sac (Text-figs. 66, 67). They are still evident during
the earlier stages of endosperm formation, but do not’ enlarge, and clearly are of
little significance. Megasporogenesis, then, and the structure of the mature
gametophyte conform to what may be designated normal, and agree very well
with what has been described in the case of Protea Lepidocarpon.

Fertilization.

The pollen tube, as already described, has been seen to enter the micropyle,
where it comes into contact with the somewhat glandular cells lining the rim
and cavity of the passage. Such cells no doubt contribute to the nourishment
of the tube, which eventually passes through the thin tissue of crushed cells of
the nucellus capping the female gametophyte, and eventually enters the embryo
sac. The discharge of the male gametes into the sac was not actually seen, but
in one preparation a crushed synergid, and two small bodies, one in the neigh-
bourhood of the egg, and another in close proximity to the two polar nuclei
(Text-fig. 69), at least suggested normal fertilization of the egg and the occur-
rence of triple fusion. These phenomena were not, however, diagnosed with
certainty.

LIFE-HISTORY OF GREVILLEA ROBUSTA,

56

BY P. BROUGH. 57

Endosperm.

The endosperm nucleus divides before that of the oospore, and subsequent
mitotic activity results in the formation of a large number of nuclei in the sac
while the egg is still quiescent, or merely increasing in size. Such early endosperm
nuclei (Text-fig. 70) are, however, practically confined to the upper two-thirds of
the sac in which region wall formation commences, and forms a tissue around
the zygote (Text-fig. 74). Meanwhile, the antipodal cells are unchanged. Subse-
quently, they neither divide nor increase in size, and so may now be dismissed,
as of no further significance. Thereafter endosperm formation proceeds slowly in
the direction of the chalaza, but is now closely pursued by the rapidly developing
embryo. The sluggish development of the endosperm in the chalazal region of
the sac is a marked feature, and even when mature this tissue often fails to fill
completely this lower end of the sac, which is often seen to be devoid of tissue,
and to collapse, when longitudinal sections of almost mature seeds are under
examination. The consequence of this behaviour, coupled with the disintegration
of the nucellus, is that the mature embryo is somewhat loosely encased by the
testa of the seed, which, owing to the complete absorption of the endosperm
during the final development of the embryo, becomes non-endospermic at maturity.

The Embryo.
; Soon after fertilization it is observed that the nucleus of the egg is consider-
ably greater in magnitude than that of the ovum from which it was derived
(Text-fig. 72). In addition, the oospore modifies its position, and moves in the
direction of the micropyle, for it is next seen in contact with the micropylar
end of the sac (Text-fig. 74). The cause of this change in position is somewhat
obscure, but one may hazard the opinion that it may be partly a natural response
lo the disintegration of the synergids, and partly due to increased pressure
consequent on the active division of the endosperm nucleus on the side of the

Text-fig. 59.—Transverse section through two impinging ovules, A and B,
of an ovary. In the centre of A is seen the large megaspore mother-cell enclosing
which are several cells more densely cytoplasmic than those more remote. The
regular radial arrangement of the cells of the nucellus is very evident. x 300.

Text-fig. 60.—Megaspore mother-cell more highly magnified than in previous
drawing. x 730.

Text-fig. 61—A megaspore tetrad. f.m., the functional megaspore adjoining
the chalaza. x 730.

Text-fig. 62.—As above, but at an older stage; all the megaspores have
enlarged. f.m., functional megaspore. x 730.

Text-fig. 63.—The functional megaspore; the other three megaspores have
been crushed and absorbed. x 730.

Text-fig. 64.—The young uni-nucleate embryo sac. Vacuolation has com-
menced. x 730.

Text-fig. 65.—Binucleate stage of embryo sac. x 730.

Text-fig. 66.—Mature embryo sac. s., synergids; o., ovum; p.n., polar nuclei
in contact; a.c., antipodal cells. x 300.

Text-fig. 67.—Lower part of embryo sac showing two of the antipodal cells,
and free endosperm nuclei. x 430.

Text-fig. 68.—Micropylar end of embryo sac with egg apparatus. s,. and s,.,
synergids; o., ovum; s.g., starch grains; v., vacuole of synergid; v,., vacuole of
embryo sac. x 730.

Text-fig. 69.—Micropylar end of embryo sac showing one normal synergid,
S,., one disintegrated synergid, S,., and ovum, O., two polar nuclei, p.7., and two
problematical male nuclei, mm. x 730.

Text-fig. 70.—Longitudinal section of embryo sac after fertilization with free
endosperm nuclei, f.e., and two antipodal cells, a.c. x 180.

58 LIFE-HISTORY OF GREVILLEA ROBUSTA,

LLG
\G

ae

su

MUTT A

tt Fl

=
Ss

Sess Sea

pS a a OS

lia

a

Text-fig. 71.—Longitudinal section cut in antero-posterior plane of ssed not
long after fertilization. o0.i., outer integument; ii., inner integument; mi.,
micropyle; n., nucellus; e.s., embryo sac; v.b., vascular bundle of funiculus, f.
x 120.

Text-fig. 72.—Nucleus of egg just before latter moves to tip of embryo sac
adjoining micropyle. x 730.

Text-fig. 73.—Longitudinal section through micropylar end of seed. The
endosperm protrudes slightly into the micropyle. o0.i., outer integument; i.1.,
inner integument; mi., micropyle; e., endosperm. x 120.

Text-fig. 74.—Longitudinal section through micropylar end of sac showing
zygote, em., amid endosperm tissue, e. x 300.

Text-fig. 75.—Two-celled stage of embryo, em., surrounded by endosperm.
x 30:0!

Text-fig. 76.—Two-celled embryo, as in previous text-fig., showing wall
separating the nuclei. x 730.

Text-figs. 77-82.—Series of figures illustrating successive stages in
embryogeny. For explanation see text. em., embryo; e., endosperm; w., wing;
o.i., outer integument; i.i., inner integument. Figs. 77, 79 x 300; fig. 78 x 200;
figs. 80-82 x 225.

BY P. BROUGH.

Text-fig. 83.—Longitudinal section of immature seed cut in lateral plane.
The embryo has differentiated into two cotyledons, c., hypocotyl, h., and
radicle, r. The wing is only shown in part, being incomplete at X and X,.
w., Wing; ii, inner integument; e., endosperm; n., remains of nucellus. x about
15. This drawing is not camera lucida.

60 LIFE-HISTORY OF GREVILLEA ROBUSTA,

egg remote from the micropyle. Certainly endosperm formation anticipates the
division of the zygote, a fact which becomes apparent on consulting Text-figure 74,
where the undivided oospore is seen embedded in endosperm tissue.

After division of the nucleus of the oospore, a wall is formed at right angles
to the long axis of the ovule. This gives the bicellular stage indicated in
Text-figures 75 and 76. Ensuing divisions lead to the formation of a flat and
approximately oval plate of cells (Text-figs. 78, 80), steadily invading the
endosperm tissue, which is being brought into solution by the secretion, of an
enzyme, and subsequently absorbed by the growing embryo. The action of such
an enzyme is amply demonstrated by the clear region fringing the embryo,
and the corroded nature of the adjacent endosperm cells (Text-figs. 78, 80, 81, 82).

At the stage depicted in Text-figure 80 periclinal walls have resulted in the
separation of a dermatogen, and inspection of successively older stages shows
the differentiation of the plerome and periblem regions, accompanied by a change
from the early oval form to one which is practically circular (Text-figs. 81, 82).

A noteworthy feature of this disc-shaped embryo is the entire absence of a
suspensor. The embryo at its inception takes up a position—at the micropylar
end of the sac—which it never vacates. Such a disposition no doubt entails
certain disadvantages from the point of view of easy nutrition, but this handicap
evidently presents no real difficulty, since further development proceeds apace,
and eventually results in the total destruction of the endosperm. During
embryogeny the embryo sac and endosperm steadily increase in volume, and some
slight invasion of the micropyle becomes manifest (Text-fig. 73).

The next significant phase is the differentiation of the main body regions—
the two cotyledons, the plumule, hypocotyl and radicle (Text-figs. 83, 84). The
last mentioned text-figure shows the adaxial surfaces of the two cotyledons, one
of which has been detached from the young plant. A noteworthy and distinctive
feature is the possession of two lobes at the base of each of the two cotyledons
which enclose the plumule and hypocotyl. A clearer perception of the nature
oi this lobing—together with the general structure of the embryo—may perhaps
be gained by examination of a mature seed in transverse section. Text-figures
86-88 show three such selected sections arranged in basipetal succession. In
Text-figure 86 the two cotyledons with their vascular bundles are apparent. The
succeeding drawing depicts the same cotyledons in the region where they are
fused with the cotyledonary zone of the young stem, which is provided with a
vascular system of four bundles.

The third section illustrates clearly the four lobes of the cotyledons. It will
be observed that they are quite detached from the hypocotyl, and that the twe
lobes on the right belong to one cotyledon, while those to the left are connected
with the other. Further, seeing that no leaves have yet grown from the plumule,
and since the stele of the young stem shows a ring of four bundles, it follows that
each cotyledon must send two separate vascular strands into the hypcotyl. Each
of these bundles branches freely at the base of the broad, flat, oval cotyledon,
as is illustrated in Text-figure 84, and so provides the vascular supply of the
seed leaves.

The Seed: Development and Structure.

Just after fertilization the seed is long, and much flattened in the antero-
posterior plane (Text-fig. 71). Consequently, in longitudinal section, its appear-
ance varies according to the plane in which the cut is made. The nucellus
is of the massive type. The inner integument is closely applied to the nucellus,

BY P. BROUGH. 61

and in turn is encased in the outer integument, but it is only at the base of
the ovule that these three components of the young seed are in organic con-
nection: they are easily separated from each other by a little pressure.

The embryo sac extends from the micropylar region of the ovule to within
a short distance of the chalaza, and in the young seed, when the embryo consists
of only a few cells, is completely enveloped in nucellar tissue, which, however,
thins out to a few layers at the upper end of the sac. Elsewhere the envelope
of nucellar tissue is relatively broad in the antero-posterior plane, especially at and
towards the chalazal end, where it forms a thick pad. Towards the periphery of
this tissue its cellular nature at this stage is still fully preserved, but internal
to this the cells have a corroded appearance which is gradually accentuated,
until, in the immediate vicinity of the sac, a cellular structure is no longer
apparent. It is surmised, therefore, that a powerful enzyme, capable of bringing
the nucellar tissue into a soluble condition, is secreted by the sac, which con-
sequently becomes suspended in a liquid nutrient medium.

As development proceeds and the endosperm and embryo become differentiated,
fresh invasions are made on the nucellus, with the result that eventually the whole
of the nucellar tissue—with the exception of the occasional occurrence of a few
remnant cells adjoining the integument and chalaza—is destroyed.

The disintegration of the nucellus is accompanied by a tremendous increase
in the size of the sac, but its increase in volume fails to keep pace with the
space vacated by the nucellus, and consequently an unoccupied region is left
between the sac and the integumentary tissue. This space is permanent, and
persists even in the mature seed. A very striking feature, consequent on
fertilization, is the very rapid and great increase in the dimensions of the seed
(Text-fig. 83). This is due, for the most part, to the tremendous activity of a
group of meristematic cells located in the basal region of the nucellus. This
meristem is apparent even in the youngest stages of the ovule, and maintains
its identity throughout. After fertilization the increase is chiefly in a direction
parallel to the long axis of the seed, which consequently is of a much elongated
and pendulous nature at maturity.

The Wing: Development and Morphology.

The mature seed is characterized by the possession of a broad, membranous
wing (Text-figs. I (27) and 71, 738, 83 and 85-88) expanded in the same plane
as the flattened body of the seed, and entirely encircling it. This structure con-
tributes to, and emphasizes the outstandingly platyspermic nature of the seed.

The morphology of this structure demands elucidation. Is it integumentary
or arilloid? In order to settle this question the development of the ovule was
studied from its inception until it passed to the mature seed condition. Such
an investigation showed that no enveloping structure, other than the integuments,
ever arises. Accordingly, the wing is a specialized integumentary development.
In the ovule the inner integument, as well as the outer, is considerably extended
in the antero-posterior plane (Text-fig. 71), and this helps to emphasize the
flattened nature of the ovule. This inner integument reaches its maximum soon
after fertilization of the ovum is effected, and is then many cells in thickness
in its broadest dimensions. Thereafter, post-fertilization growth of the tissues
within, and resistance offered by the enveloping integument without, result in
lateral crushing which is accentuated until, in the mature seed, only a thin,
almost structureless tissue represents a formerly outstanding component of the
ovule.

62

LIFE-HISTORY OF GREVILLEA ROBUSTA,

rf
Ht
tH
Hh
HH
te
a
Ho
HA
sa
Hy
8)
Ht
ta]
th
Hh
9
ta

aS

Vatu
SOausaaee

oe

Text-fig. 84.—Embryo removed from seed. The cotyledon on the right
has been detached from the young plant. The adaxial surfaces of the cotyledons
are exposed. opl., plumule; h., hypocotyl; r., radicle; v.b., vascular bundles;
l., lobe of cotyledon. x 30.

Text-fig. 85.—As in Text-fig. 83, but on a higher scale of magnification and
with the central portion of the seed omitted. em., embryo; w., wing; v.b.,
vascular bundle. x 30.

Text-figs. 86-88.—Selected sections from a series of transverse sections of
mature seed. For explanation see text. Only in Text-fig. 87 is the full extent
of the wing shown. w., wing; l1., lobes of cotyledon; ¢., cotyledonary zone;
h., hypocotyl. x 30.

Text-fig. §9.—Longitudinal section, cut in lateral plane, through micropylar
end of seed showing detailed structure of part of wing. o0.i., outer integument;
ii., inner integument. x 110.

BY P. BROUGH. 63

In the case of the outer integument, however, a very different fate falls
to be recorded.

At the stage of ovule development illustrated in the text-figure last mentioned,
its structure is very similar to that of the inner integument, but soon after
fertilization, and at the stage when the endosperm has become conspicuous (Text-
fig. 73), meristematic activity is noticeable in the row of cells next to those
forming the inner limit of the integument. These divide rapidly and in orderly
fashion, and thus form a much extended tissue. The radial arrangement of
the cells resulting from this regular tangential division becomes more noticeable
as development proceeds (Text-fig. 89). This lateral expansion of the integument
continues, however, to be much more active in the antero-posterior plane, with
the result that the integument assumes the flattened form characteristic of the
wing in the mature seed.

The lateral involutions seen in the transverse section of the seed (Text-
fig. 87) are due to the outer integument accommodating itself during later
development to the original lateral expansion of the inner integument. Such
expansion of both integuments in the antero-posterior plane is, as already stated,
a feature of the ovule just prior to fertilization, but whereas the outer integument
continues to increase in size, the inner integument suffers compression and
practical obliteration, but none the less records its previous existence in the
lateral involutions evident on each side of the mature seed when viewed in
transverse section.

It must be emphasized that a very marked feature of the development of the
seed as a whole is a great increase in length accompanied by excessive develop-
ment in the antero-posterior plane. This latter feature is an outstanding char-
acteristic of the development of each component part—nucellus, endosperm, embryo,
integuments—of the ovule and seed, and accounts for its mature form. Accord-
ingly, the pronounced flattened shape of this organ and the morphological nature
of the wing have now been interpreted.

Pollination.

The anthers dehisce just prior to or simultaneously with the process of
anthesis, so that when the style frees itself from the perianth segments, the
pollen is poised as a cone-shaped mass on the distal end (Text-fig. I, 10). This
occurs before the stigma is exposed. Accordingly pollen is prevented from coming
into contact with the stigma of the flower which produces it. In other words,
the flower is strictly protandrous. Practically all of this pollen is dislodged
during the first day or two of its exposure, either by the numerous bird visitors,
or by air currents. In any case, in the event of any pollen retaining its original
position, it is certain to be dislodged—outwards and downwards—when at a
later stage the growing stigmatic cells finally burst through the epidermal layer
(Text-fig. 34) and first expose themselves to outside contact. The whole process
is very much akin to the procedure during the pollination period of Dampiera
stricta (Brough, 1927, p. 493).

On further development a wider crater-like cavity, which is lined by the
long papillate cells of the stigma, is formed (Text-figs. 37 and 38). Some of these
receptive cells protrude from the opening, and are in a suitable position for
the reception of pollen. Owing to its general configuration and the nature oi the
cells composing it, such a stigma is eminently suited to the germination, nourish-
ment and development of the young male gametophyte.

64 LIFE-HISTORY OF GREVILLEA ROBUSTA,

Below the stigma proper is located a tract of thin-walled elongated cells
(Text-figs. 33 and 37) which traverses the length of the style, and provides an
admirable conducting and feeding tissue for the passage of the pollen tube.

In the section of this research dealing with organogeny and general floral
development, attention has already been drawn to certain abnormal flowers in
which the perianth is not shed at the usual time, but persists as a cap enveloping
the distal end of the style (Text-fig. I, a and Bp). One naturally wonders concern-
ing the fate of such “capped” flowers, and the question arises is self-pollination
effected, and if so, do embryos arise?

It so happened that the answer to such queries was suggested during the
examination of another species of Grevillea, namely, G. Banksii, a species in which
“capped” flowers (Text-fig. I, a-f) were found to be much more abundant than
in the Silky Oak. Incidentally, it may be mentioned that observation showed that
in G. Banksii seeds are set more abundantly than in any other species known
to the writer. Here, in one selected case, 25 “capped” flowers, out of a total of
42 on a single inflorescence, were found to have full-sized fertile seeds, although
the perianth still protected the stigma. This showed quite conclusively that self-
pollination and fertilization were by no means uncommon in this species. In
fact, from observations made it is computed that during the latter part of the
seed-setting season thirty to forty per cent. of the seeds produced in this district
by G. Banksii are derived from “capped” flowers. <A series of photographs illus-
trating successive phases in the development of a “capped” flower from the bud
stage till the period of full seed development is depicted in Text-figure I, a—f. Just
how prevalent this phenomenon is in other species of Grevillea and throughout
Protead forms in general is at present unknown, but the author hopes to supply
information on this point in the near future.

Possessed of a knowledge of the significance of “capped” flowers in G. Banksii,
the writer decided to prosecute a similar investigation in the case of the Silky
Oak. A survey of various trees revealed the fact that “capped” flowers, though
occurring occasionally, are much more rare than in the species already described.
However, specimens were collected in which the corolla persisted long after the
perianth segments had been shed from the remaining flowers of the same or
adjoining inflorescences. In two selected inflorescences gathered on 3rd November,
1931, one raceme of 65 flowers had 6 “capped” individuals, while in the other
with 81 flowers, the number of ‘capped’ specimens was 13, i.e., an average of 14
per cent. This ‘‘capped’’ condition then does occur, though sparingly, and persists
beyond the period of normal pollination, and in such cases the increase in size
of the ovary clearly points to the fact that in this species also self-pollination
and seed-setting do take place. The question then arose as to why the phenomenon
is so much more restricted in the case of Grevillea robusta than it is in Grevillea
Banksii. Examination revealed that in the former species the perianth segments
are much more easily ruptured at the base and apex, and that the repeated
contact of numerous birds—large and small—in their search for nectar expedites
the rupture and fall of the perianth.

It is only in special cases then—probably where a flower is relatively
inaccessible, as at or near the tip of an inflorescence, or where bird visitors for
some reason are rare—that the perianth may not be dislodged. The important
points that emerge, however, are that should bird visitation fail, then the number
of “capped” flowers will be increased, the chances of self-pollination be enhanced,
and accordingly a compensating factor operates for the setting of seed.

BY P. BROUGH. 65

Further investigations during this season concerning the incidence and
significance of “capped” flowers in this species were unfortunately rendered
nugatory by the advent of an exceedingly severe windstorm which, by displacing
the perianth segments, destroyed all evidence as to “capping” tendencies.

Having demonstrated the structure of the flower about the time of anthesis
and pollination, it is natural to transfer one’s attention to the subject of pollinating
agents.

Observations taken during a clear, sunny day in mid-November, 1930, between
the hours of 10 a.m. and 4 p.m., on a tree about 60 ft. high at Wahroonga, some
twelve miles from Sydney, revealed the presence of numerous birds of various
kinds. These visitors were actively at work sipping the profuse nectar, and at no
period of the day was this particular tree devoid of feathered species of some kind.

The birds noticed in greatest number were several species o: honeyeaters,
and it was noteworthy that one of the less common species, viz., Meliphaga
melanops (the Yellow-tufted Honey-eater) was present in considerable numbers.
Also abundant were Meliphaga chrysops (the Yellow-faced Honey-eater) and
Acanthorhynchus tenuirostris (the Spinebill Honey-eater). In addition, Antho-
chaera chrysoptera (the Little or Brush Wattle Bird) and Philemon corniculatus
(the Noisy Friar Bird or Leather Head) formed a conspicuous component of the
bird population. Other birds were occasionally seen, but those above mentioned
were regarded as the forms responsible for the removal of nectar from the flowers
of the Silky Oak.

The force exerted by the beak of the bird, while probing for nectar, causes
the style to bend forward and downward, thus bringing the tip into contact with
some part of the bird’s head, which is accordingly dusted with pollen in the
case of flowers recently opened, or is brushed over by the stigma in more mature
flowers. In this way cross-pollination is readily effected, since a single bird was
seen to visit many flowers within a few minutes.

It is worthy of note that the pollination mechanism in Grevillea Banksii
is much more exact than in Grevillea robusta, for in this case the apex of the
style and stigma are flattened, and the flower so constructed that the stigmatic
disc always comes down on the same spot on top of the visitor’s head. This clearly
increases the chances of pollination and, taken in conjunction with the close and
frequent association of Meliphaga melanops with this shrub, would seem to be an
important factor in accounting for the relatively high percentage of seed set in
this species of Grevillea.

In addition to birds, however, the writer constantly observed insects—chiefly
bees and ants—in great numbers at work on the flowers of the Silky Oak. These
move about at the bases of the flowers, passing from one to another, but, after
the freeing of the style and consequent exposure of the pollen, the dimensions
of the flower are such that the stigma is too far removed from the nectar to
permit of contact between the visiting insect and the stigmatic region, unless
perhaps in cases of mere chance. Such visitors, then, are mere nectar-robbers
and in no way associated with pollination. On the other hand, birds, on account
of their larger size, are pollinating agents, and so Grevillea robusta and Grevillea
Banksii are strictly ornithophilous forms. However, in species of Grevillea
with smaller flowers, e.g., Grevillea sericea, bees were observed to be the pollinating
agents.

The infertility of the Proteaceae in general has long been known, and widely
commented upon.

H

66 LIFE-HISTORY OF GREVILLEA ROBUSTA,

Within recent years Lawson (1930, p. 374) has drawn attention to the very
high percentage of sterile pollen in numerous Protead forms investigated by him,
and supplies in tabulated form a list of figures giving actual percentages. Therein
Grevillea robusta is quoted as having ninety per cent. of sterile pollen, and
this fact alone might perhaps satisfactorily account for the relatively low seed
production.

Actuated by impressions gained from a study of sections of anthers, which
suggested that the figures given by Lawson in the case of Grevillea robusta were
unduly high, the writer made fresh estimates. Lawson’s methods were adopted,
although such seem only capable of supplying approximate results. In no case
was the percentage of sterile pollen found to exceed 30 per cent., while the figure
for Grevillea Banksii was in the neighbourhood of 20 per cent. Again, reference
to the photograph of the microspores of Grevillea robusta accompanying Lawson’s
paper (Lawson, 1930, Plate x, B) hardly seems to support the high percentage
claimed.

Accordingly the writer does not subscribe to the opinion that pollen sterility
is the dominating factor in accounting for low seed production, although it is
one of the contributing factors. In this connection it is interesting to observe
that Hamilton (1931, p. xl) in reviewing Lawson’s paper, concludes with the
following statement: “I think, therefore, that a good case has been made for
the theory that the sterility of the Proteaceae is caused, to a great extent, by
the failure of the pollinators to visit flowers, and that it is not due to any
inherent infertility.”

In the case of Grevillea robusta, nectar is produced in such quantity that,
on shaking a branch, drops shower down in sufficient quantity to cause unpleasant
staining of one’s clothes, while handling an inflorescence roughly results in the
smearing of the hands.

Nectar is present even before the style is freed, and so at this stage its
removal can be of no direct aid in pollination, although the movement of the
birds undoubtedly accelerates actual anthesis.

From the facts put forward it would seem, then, that in the case of the
Silky Oak, and probably other Protead forms as well, the chief factor control-
ling seed setting is the visitation of pollinating agents, although the results are
modified to some extent by the relatively high percentage of sterile pollen. This
latter, however, is to be regarded as a minor factor.

But the matter does not end here; another problem arises: Why is there
such a great variation in the amount of seed produced from year to year in
Silky Oaks generally, and also in the individual tree? For example, the seed
production in 1931 was abnormally high, while the present season’s crop for
1932 is exceedingly meagre. Two reasons underlie this phenomenon: first, the
exposed position of the pollen, and second, the weather conditions prevailing
during anthesis of any one plant, and of Silky Oaks in general.

In 1930, during the period of maximum anthesis and pollen exposure, the
climatic conditions were uniformly dry and sunny without acute atmospheric
disturbances. Such conditions presented an ideal environment for pollination,
and consequently a high seed production was recorded in the early part of 1931.

But during the pollination period of 1931, abnormally dull, cold and wet
weather prevailed, and a wind storm of intense violence swept the neighbour-
hood. In other words, the weather was most unseasonable in regard to low
temperatures, high winds, and heavy rains.

BY P. BROUGH. 67

Such conditions naturally were antagonistic to pollination on account of
general damage to flowers, but more particularly to the displacement of the
pollen, which was scattered and lost, and also to the very greatly increased
percentage of sterile pollen which had been adversely affected by repeated wetting
and drying. During this season, then, pollen sterility was a big factor in
accounting for low seed production, but this sterility was due to secondary
causes, and not inherent in the original pollen as produced by the flowers.

Different plants open their flowers at different times throughout the general
period of anthesis, and this fact, coupled. with whether the individual plant
occupies an exposed or sheltered position, accounts for the vagaries of seed
production from tree to tree, during any particular season.

A résumé of the facts submitted concerning pollination and fertilization of
Grevillea robusta urges the writer to the conclusion that seed-setting from year
to year is controlled by two main factors, the relative abundance of pollination
visitors and the meteorological conditions prevailing during the chief period of
pollen exposure. If either of these factors is unfavourable the seed production
will be low, while if both are inimical to pollination then very few seeds indeed
will be produced. On the other hand, if both factors act simultaneously in
favour of reproduction, then a maximum seed crop will result.

It is suggested that the conditions controlling seed production in Grevillea
robusta are of general application in the Proteaceae.

Comparison with Protea Lepidocarpon.

Having obtained a knowledge of the salient features in the life-history of
Grevillea robusta, one naturally proceeds to effect a comparison with corres-
ponding phases, so far as they are available, in Ballantine’s account (1909) of
Protea Lepidocarpon. Generally speaking the correspondence is very close, and
marked agreement is noted in the pendulous nature of the ovule; the single
megaspore mother-cell situated amid a group of large cells below the hypodermal
layer; the definite meristematic tissue at the base of the nucellus which remains
active until time of fertilization; the linear tetrad of megaspores, the chalazal
one of which functions; the shape of the embryo sac, and its protrusion, though
slight, into the micropyle; the glandular nature of the cells at the tip of the
nucellus, and also of the cells of the inner integument surrounding the micropyle;
the disappearance of the nucellar tissue soon after fertilization; the absence of
a suspensor in, and the globular form of, the embryo and the method of develop-
ment of the microspores and male gametophyte.

On the other hand, certain contrasting features are to be noted, the most
outstanding of which are that the haploid number of chromosomes is only ten,
that no irregular protuberances of the nucellus into the integuments are present,
that the origin of the ovules is in the middle region of the ovarian cavity, and
that the embryo sac does not become densely packed with starch, although
starch grains do aggregate in quantity in the neighbourhood of the ovum just
prior to ‘fertilization.

These dissimilarities, however, do not to any significant extent detract
from the essential agreement of the phases available for comparison, or from
the validity of the deduction that a very close affinity exists between the two
genera under comparison.

Conclusions.
* A study of the floral organogeny shows that the perianth is essentially
primitive, consisting, as it does, of four free segments arising from the rim of

68 LIFE-HISTORY OF GREVILLEA ROBUSTA,

a very slightly concave torus, which in turn is slightly inclined to the plane at
right angles to the axis of the flower. There is no evidence that there ever has
been any differentiation into a calyx and a corolla, or that the flower represents
a reduced type.

The flower has bilateral symmetry, the plane being the antero-posterior one.
Zygomorphy is induced by the later development of the style, ovary and ovules
being accentuated in the plane referred to. The early fusion of the meristematic
zones, which give rise to the perianth segments and stamens respectively, is a
feature of advance, but one easily effected. Moreover, the degree of fusion varies
throughout the Family. In fine, the facts of organogeny relative to the perianth
and androecium indicate an essentially primitive structure for the flower in
question.

The development of the anther is characterized by the production of a
thick wall, a fibrous layer of hypodermal origin, a plate one cell thick of spore-
mother-cells, a normal tapetum, and a somewhat unusual, but none the less
simple, method of dehiscence.

The carpel shows very clearly its single, foliar origin, and bears two ovules
with marginal placentation. These ovules are anatropous, but as anatropy is
a feature of early evolutionary attainment, the gynoecium as a whole, and the
resulting fruit, are essentially primitive in structure. Simplicity in fruit
structure is characteristic of the family.

Pollination in the Proteaceae is effected either by insects or birds, and is
also associated with self-pollination, but the mechanisms involved are not of a
type embodying any high degree of specialization in floral structure.

Again, an appreciation of the salient features in the life-history of Grevillea
robusta brings a realization of the marked simplicity of the type chosen for this
study. Further, as already explained, a survey of the outstanding phases in the
life-histories of Grevillea robusta and Protea Lepidocarpon results in the realiza-
tion that a very close agreement exists between the two forms, but the chief
interest of such a comparison is centred in the fact that certain peculiarities in
Protea Lepidocarpon, such as the glandular cells limiting the apex of the
nucellus, the similar cells associated with the micropyle, the suggestion of a
former multicellular sporogenous tissue in the ovule, and the rounded embryo
devoid of a suspensor, are all represented in Grevillea robusta.

The obvious deduction is that such peculiarities could not have been evolved
independently, but have been inherited from a common ancestry. Now, present
land distribution does not permit of direct contact between Africa and Australia;
moreover, the separation of these two continents seems to have persisted since
Carboniferous times when Gondwanaland connected Australia, India and South
Africa (Berry, 1918).

Again, Australia is considered to have been joined with the Asiatic main-
land at least during the upper Cretaceous period (Benson, 1923, pp. 50-52), but
probably a complete separation has existed since the beginning of the Hocene.
From such facts one may infer that the ancestors of the Proteaceae—as well as
those of other Australian Families—migrated southwards from an Asiatic centre
of distribution into Australasia and Africa, as then constituted, during later
Cretaceous times. It may be stated that an intensive study of the Epacridaceae
by the writer fully supports the above contention.

Keeping in mind the general characters of the Family as a whole, and of
the details regarding Grevillea robusta made available as a result of this

BY P. BROUGH. 69

investigation, and of Protea Lepidocarpon as described by Ballantine, one may
proceed to a consideration of the affinities of the Proteaceae and the real position
of this Family in a natural scheme of classification. As already pointed out
in the early pages of this publication, different writers have claimed close
kinship for the Proteaceae with the Santalales, Thymeleaceae, and Rosales
respectively.

At this juncture the writer believes that little purpose would be served in
recounting and reconsidering the minutiae of detail pertaining to gross
morphology. This type of evidence has already been marshalled and dealt with
by eminent systematists, and with the results indicated. Further progress is
rather to be achieved by comparing details of the life histories—so far as they
are available—of the forms most intimately concerned with the purpose in view,
gross characters only being used as a check on deductions made from other
evidence,

Dealing with the Santalales claim, it may be said that the evidence from
gross morphology is not of the most convincing nature. If any real affinity does
exist, then the modifications which have taken place in the various Santalalean
Families since the common connecting type was departed from, are so great that
features indicative of any real kinship have become obscure, almost to the
point of obliteration.

Turning to the evidence supplied from investigations relative to life-histories,
much information has been supplied by Guignard (1885), but comparison with
corresponding phases in Grevillea robusta fails to substantiate any claim for
close kinship. Again, in more recent times, Skottsberg (1913) has investigated
the morphology and embryology of the Myzodendraceae, but here again the
results are negative, for although the embryology runs parallel with that of
the early and medium stages in Grevillea robusta, there is nothing beyond that
on which any near relationship may be based.

Turning now to the claim for a Proteales-Thymeliales affinity, it may be
conceded that, despite considerable variation in floral structure within the
Families of the Thymeliales (Hutchinson, pp. 150-152) there is nothing in gross
morphology really incompatible in such an association. A nice series in the
gradual fusion of the perianth segments is illustrated in passing from the
Proteaceae where fusion is suggested, to the Geissolomataceae where the “calyx”
is shortly tubular and four-partite almost to the base, and thence to the
Thymeleaceae and Penaeaceae where the “calyx” is markedly tubular. Throughout,
the stamens are epiphyllous and the ovary superior. Accordingly a more
searching scrutiny may be the more hopefully instigated. An approach to the
problem may best be made by reference to the investigations of Stephens on the
Penaeaceae (1908 and 1909) and also on Geissoloma marginatum (1909a). Dealing
first with the Penaeaceae, it is to be noted that in the three genera examined,
viz., Sarcocolla, Penaea and Brachysiphon, the development of the female
gametophyte is abnormal, Stephens (1909c, p. 381) pointing out that “the first
two divisions form a tetrad of nuclei arranged just as in Peperomia: these
separate and each divides again twice, so that four groups of four nuclei each
are formed at the periphery of the sac. Cells are organised round three of the
nuclei in each group, in the form of an egg-apparatus.”

There is, of course, nothing like this in Grevillea robusta, and so far there
is no general agreement as to the significance of, or cause underlying, the
abnormal behaviour noted in the Penaeaceae and Peperomia. Further, Stephens

70 LIFE-HISTORY OF GREVILLEA ROBUSTA,

(1909, p. 370) says: “As regards the other Orders with which the Penaeaceae
are grouped, the Thymeleaceae is the only one of whose life-history any account
has been published, Strasburger having recently investigated species of
Wiksstroemia, Daphne and Gnidea, none of which show a similar departure
from the normal.”

The embryology of the Penaeaceae conforms closely with that of Geissoloma
marginatum (Stephens, 1909a), but it would not be valid to establish any connec-
tion on that feature alone. Accordingly, the Penaeaceae, if retained within the
Order Thymeliales, must be regarded as occupying an isolated position, and with
its kinship hon proven.

However, on comparing Geissoloma marginatum (Stephens, 1909a) with
Grevillea robusta marked similarities are evident in the general structure of the
nucellus, the normal development of the functional megaspore and female
gametophyte, the embryo—in the early stages oval, and later becoming round—
the emergence of the cotyledons amid a greatly elongated copious endosperm,
which in turn lies in a sap-containing cavity due presumably to the collapse of
the nucellus. Accordingly, the evidence justifies one in recognizing an affinity
between the Proteaceae and the Geissolomataceae without admitting any close
connection between these two Families and the Penaeaceae. At such a stage one
feels how unfortunate it is that the investigations referred to were not of a
wider scope in the individual, so that other points of contact might be sought.

Finally, there remains the claim that the Proteales may best be associated with
the Rosales. The amount of ground which would of necessity be covered in
making a critical examination of such a claim would be so extensive as to extend
impracticably the scope of this section of the present investigation. Moreover,
amid the plexus of forms included in this Order there is no obvious section to
which the inquiry might be confined, and more important still is the fact that
the writer is convinced that much more research must necessarily be effected in
ihe realm of adjoining and comparatively unknown Orders and Families before
such a task could be attempted with any hope of success commensurate with the
labour involved.

Suffice it to say in conclusion that, while the results of this research on
Grevillea robusta are not incompatible with a real affinity between the Proteaceae
and the Geissolomataceae and through them with the Thymeleaceae in general,
still the view must not be lost sight of that the simplicity of the Proteaceae may
well be primitive, in which case an affinity with some Order much lower in the
scale of evolution than the Thymeliales will have to be sought. The success
of such a quest will probably depend on the extension of such investigations as
the one now completed.

SUMMARY.

Floral Organogeny.—The primordia of the individual members of the various
sets of floral organs arise separately, and in acropetal succession. The torus is
very slightly concave and from its rim originate the four separate primordia of
the perianth segments, which later simulate gamophylly owing to the interlocking
of the contiguous marginal cells. There is no indication that a separate calyx
and corolla ever existed. The stamens arise as individuals from the torus, but
become epiphyllous at a very early stage. A single carpel occupies the organic
apex of the floral receptacle, and soon after its inception, a crescent shaped nectar-
scale arises between the gynoecium and the perianth segments, and on the adaxial
side of the flower.

BY P. BROUGH. 71

Microsporangium.—The archesporium consists of a plate one cell thick
which divides by periclinal walls giving rise to a primary parietal and a primary
sporogenous layer respectively. The cells of the former divide repeatedly, thereby
producing a wall four cells thick lined internally by a tapetum forming a fifth
layer. The sporogenous cells, without further division, function as the spore
mother-cells, which undergo reduction division, whereby pollen tetrads are
produced. At dehiscence the microspores are binucleate, and tetrahedral in shape,
with a pore at each of the four corners. The exine is very thick and heavily
cutinized. The fibrous layer is hypodermal, and dehiscence is effected by the
orderly separation without rupture of the overlapping margins of adjacent micro-
sporangia—a somewhat rare mechanism in Angiosperms.

The male gametophyte—The mature microspores contain a small generative
and a larger vegetative nucleus separated by a thin membrane. Just prior to
germination vacuolation becomes evident, and a single pollen tube emerges at
one of the four unthickened areas of the pollen grain. Germinating grains were
seen on the stigma, and the pollen tube was traced in its growth down through
the style, and along the wall of the ovarian cavity, whence it crossed to the
ovule near the apex and, growing along its surface for a short distance, eventually
entered the micropyle, which is lined by cells of a glandular nature.

Nectar secretion.—The morphological nature of the nectar-scale is explained,
and profuse nectar secretion from this organ, and also from the cells of the
inner epidermal layer of the basal regions of the perianth segments, is described.

Ovule-——The two nucellar primordia arise from opposed margins of the
carpel, and in the region adjoining the base of the ovarian cavity.

A marked feature is that development of the ovule is almost entirely due to
the pronounced activity of a meristematic zone of cells in the basal region of
the nucellus.

An inner and an outer integument arise relatively late and develop slowly,
but eventually enclose the nucellus except in the region of the micropyle. The
nucellus is of the massive type, and eventually a single deep-seated megaspore
mother-cell is differentiated. Contiguous cells show sporogenous tendencies, but
never develop into spore mother-cells. A linear tetrad is produced, the inner-
most megaspore of which enlarges and destroys the others.

The female gametophyte.—The functional megaspore initiates germination by
vacuolation and an increase in size. This is succeeded by the bi-nucleate, four-
nucleate and eight-nucleate stages respectively of the embryo sac. The egg
apparatus, polar nuclei and antipodal cells are normal in structure and polarity.
During this development only part of the encasing nucellar tissue is invaded,
except at the micropylar end, where destruction of the nucellus is practically
complete.

Fertilization—The available evidence indicates that one male nucleus
fertilizes the oosphere, while the other joins the two polar nuclei which were
seen to fuse in close proximity to the ovum.

Endosperm.—The endosperm nucleus divides before that of the oospore
which passes to, and impinges upon, the micropylar end of the sac. Numerous
free endosperm nuclei are formed, but few of these are located at the chalazal
end of the sac. Wall formation in the upper and central regions accompanied by
a rapid extension in length of the embryo sac then supervenes. The growth of
the endosperm tissue is relatively slow, and barely fills the antipodal end of

72 LIFE-HISTORY OF GREVILLEA ROBUSTA,

the large sac. Enzyme action brings into solution, and completely destroys, the
remaining nucellar tissue.

Antipodal Cells——These remain quiescent, and have no apparent functional
significance.

Embryology.—Development of the oospore begins relatively late, but once
started proceeds with great vigour. There is no suspensor. In its early stages
the embryo is roughly oval in shape, and flattened in the antero-posterior plane.
Later it becomes round and disc-like in form and subsequently dermatogen,
periblem, plerome, cotyledons, plumule and radicle are differentiated. Hach
cotyledon has a peculiar double lobe at its base. The endosperm is eventually
destroyed.

Pollination.—Caretul attention was given to this problem. The flower is
markedly protandrous, and transfer of pollen is effected by various birds, the
names of which are specified. The pollination mechanism is described. Many
visiting insects are mere nectar-robbers. Self-pollination is possible and does
occasionally occur in Grevillea robusta. In Grevillea Banksii, however, thirty to
forty per cent. of the flowers may be self-pollinated. The phenomenon of self-
pollination may be widespread throughout the Proteaceae.

Seed structure.—The mature seed is non-endospermic and bears a broad wing
which is morphologically an extension of the outer integument.

Seed production.—The reasons underlying the variation in the heaviness of
the seed crop from year to year, and among individual trees, are given. Factors
responsible for the notoriously low seed production in the Family as a whole are
described.

Comparisons and affinities —Comparison with the South African form Protea
Lepidocarpon is made, the genetic relationships of the Proteaceae are discussed,
and some general deductions regarding plant distribution and area of origin
of the Family are recorded.

Acknowledgments.

The writer desires to acknowledge his indebtedness to Mr. H. C. Barry,
Wahroonga, for assistance in identifying the birds associated with pollination,
and to thank Mr. O. D. Evans, Laboratory Assistant, Botany Department,
University of Sydney, for valuable aid in procuring the material for this
investigation,

List of References.
BALLANTINE, A. J., 1909.—Preliminary Note on the Embryo-sac of Protea Lepidocarpon.

Ann. Bot., xxii, pp. 161-162.

BENTHAM, J., 1870.—Flora Australiensis.
BENSON, W. N., 1923.—Palaeozoic and Mesozoic Seas in Australasia. Trans. New

Zealand Institute, 54, pp. 1-62.

Berry, HE. W., 1918.—Annual Report of the Smithsonian Institution.
Bews, J. W., 1927.—New Phytologist, xxvi, Nos. 1 and 2.
BroucH, P., 1923.—Preliminary Note on the Embryo Sac of Styphelia longifolia. Proc.

LINN. Soc. N.S.W., xlviii, Pt. iv.

, 1924.—Studies in the Hpacridaceae. I. The Life-history of Styphelia longi-

folia. Proc. Linn. Soc. N.S.W., xlix, Pt. ii.

, 1927.—Studies in the Goodeniaceae. I. The Life-history of Dampiera stricta.

Proc. LINN. Soc. N.S.W., lii, Pt. iv.

CAMPBELL, D. H., 1897.—A morphological Study of Naias and Zannichellia. Proc.

California Acad. Sci.

FRANCIS, W. D., 1929.—Australian Rain-Forest Trees.
GUIGNARD, L., 1885.—Observations sur les Santalacées. Ann. Sci. Nat., Bot. vii, 2, pp.

181-202.

Proc, Linn. Soc, N.S.W., 1938. PLATE

1.—Grevillea robusta. 2.—G. Banksii.

-~]
a)

BY P. BROUGH.

HAMILTON, A. G., 19381.—Note on the Sterility in the Proteaceae. Proc. LINN. Soc.
INESHVWige iv GL980)),, et. 6:

HuTCHINSON, J., 1926.—The Families of Flowering Plants. MacMillan & Co., Ltd.

Lawson, A. A., 1930.—Endemism in the Angiosperm Flora of Australia. Proc, Linn.
Soc. N.S.W., lv, Pt. 4.

MAIDEN, J. H., 1917.—Forestry Handbook. Pt. ii.

Moorr AND BEtTcHR, 1893.—Handbook of the Flora of N.S.W.

NawmMan, I. V., 1928.—Life-history of Doryanthes excelsa. I. Proc. LINN. Soc. N.S.W.,
Litinee tabs

————,, 1929.—_Life-history of Doryanthes excelsa. II. Proc. LINN. Soc. N.S.W., liv,

Pta4:
OsBORN, T. G. B., 1932.—The Plant in Relation to Water. Proc. LINN. Soc. N.S.W.,
Ivii, Pts. 1-2.

PEARSON, H. H. W., 1929.—Gnetales. Cambridge University Press.
RENDLE, A. B., 1925.—The Classification of Flowering Plants, Vol. II. Cambridge
University Press.
SARGENT, O. H., 1930.—xXerophytes and Xerophily, with special reference to Protead
Distribution. Proc. Linn. Soc. N.S.W., lv, Pt. 5, pp. 577-578.
SKOTTSBERG, C., 1913.—Morphologische und Embryologische Studien uber des Myzoden-
draceen. Kung. Sven. Vet. Ak., Hand., Bd. 51.
STEPHENS, E. L., 1908.—A Preliminary Note on the Embryo-sac of certain Penaeaceae.
Ann. Bot., xxii, pp. 329-330.
,1909a.—The Embryo-sac of Geissoloma marginata. New Phytologist, viii, pp.
345-347.
,1909b.—The Embryo-sac and Embryo of certain Penaeaceae. Ann. Bot.,
Xxili, pp. 363-378.
————,, 1909ce.—Recent Progress in the study of the Embryo-sac of the Angiosperms.
New Phytologist, viii, p. 381.

EXPLANATION OF PLATE II.

Fig. 1.—Foliage and inflorescence of Grevillea robusta. The individual racemes of
the paniculate inflorescence are apparent, while the fact that the flowers arise in pairs
may be observed. x 4.

Fig. 2.—Raceme of Grevillea Banksii, demonstrating the formation of fruits despite
the fact that the stigma is still completely enclosed within the “cap’’ formed by the
cohesion of the distal regions of the perianth segments, the bases of which have been
detached from the floral receptacle. The phenomenon is not uncommon in this species,
and shows that self-pollination is of frequent occurrence. x 3.

NOTES ON AUSTRALIAN DIPTERA. XXXIII.
By J. R. Mattocw.
(Communicated by F. H. Taylor.)

(One Text-figure. )
[Read 26th April, 1933.]

In this short paper I present a synopsis of the species of the genus Amenia
Kobineau-Desvoidy, belonging to the Ameniinae, and the description of one new
genus of the same Family, Tachinidae, which is referable to what is generally
accepted as a distinct tribe under the name Trichopodini, a group which up to
the present time has not been recorded as occurring in Australia.

I regret that because of much miscellaneous material being still in my hands
in an unidentified condition it is impossible for me to publish a more extensive
key to the Australian genera, as several genera of uncertain status still remain
to be elucidated. I hope, however, to complete my work within the year and will
then publish a full generic key.

At the time the following notes on the genus Amenia were written there
seemed to be little chance that additional data would be obtained upon the
Robineau-Desvoidy collection, but I have subsequently been informed that at
least a part of this collection has been found, though to what extent it may
prove of service in the elucidation of the Australian species described by that
author, or indeed if it contains the types of any such species, is unknown to me
at this time. There does not appear, however, to be any likelihood of a change
in the status of the only two species involved, Amenia imperialis Robineau-
Desvoidy and A. leonina Fabricius, so I present the data as available to me.

Genus AmENIA Robineau-Desvoidy.

This genus contains several very beautiful species, all of which are confined
to Australia. There has been some difference of opinion as to the synonymy of two
of the older described species, leonina Fabricius and imperialis Robineau-Desvoidy,
and it is my intention to point out briefly herein the reason why I have arrived
at my conclusion that the two names mentioned above are properly applicable to
distinet species.

AMENIA LEONINA (Fabricius).

Syst. Hnt., 1775, 776; Hnt. Syst., iv, 1794, 318; Malloch, Proc. Linn. Soc.
INES cg Wy Dey 24 AUDETOS Ital

Wiedemann redescribed this species from the material in the Fabricius
collection and in his description he specifically mentions the silvery anterior
vittae on the mesonotum, ‘“‘Ruckenschild erzgrun, vorn mit vier wenig merklichen,
kupferrothlichen striemen, deren mittlere in gewisser richtung silberschimmern
(lineola apicis des Fabricischen textes)’’, and the description of the head contains
the following that applies to only the specimens that I have determined as
belonging here: ‘“Hinterkopf duster erzgrun, mit rothlichem, den_ scheitel

BY J. R. MALLOCH. 75

erreichenden mittelflecke.’ The abdomen, he states, has ‘an jeder seite des
zweiten abschnittes steht ein silbertropsen nahe am seitenrande und ein anderer
minderer sichtbarer, ja in gewisser richtung ganzlich verschwindender, in der
mitte jeder seite’.

AMENIA IMPERIALIS Robineau-Desvoidy.

Essai sur Myod., 1830, 443; Malloch, Proc. Linn. Soc. N.S.W., 55, pt. 2, 1930, 101.

The author of the second species had before him what he accepted as two
species; one he considered to be leonina and the other he described as new under
the above name. He gives the description of the thorax as follows: “corslet d’un
beau vert-dore metallique, avec trois points argentes de chaque cote du dos”, and
the abdomen ‘‘d’un beau vert-dore metallique, avec deux petits points blancs sur
les cotes du second segment et deux autres points blancs plus larges sur le
penultiéme segment’’.

His distinction between the two species under the heading of the older one
is as follows: “Elle différe de 1’A. imperialis par ses teintes azurées, par sa face
ferrugineuse, par la presence de deux petites lignes argentées vers le sommet de
l‘écusson, et par ses cuillerons moins blancs.”

There are unquestionably two species here, the males being very distinct in
that the frons in leonina is reduced to a mere line on its upper portion, while
in imperialis the male has the frons about as wide as or wider than the length
of the antenna. In the females the frons is narrower in leonina than in imperialis
also, and in both sexes the dark green upper outer halves of the back of the
head are conspicuously different from the densely golden-yellow dusted upper
halt of the occiput in imperialis.

I have never been abie to detect any fine hairs on the membranous strip
below the lower calypter in imperialis, but in many specimens of leonina there
are such hairs, though the feature is not constant enough to justify the use of
it as a specific character.

In connection with the above notes it may be of interest to note that Dr.
Engel, in his paper in 1925 (Zool. Jahrb., 50, pp. 350-353), uses the name Jeonina
for what I accept as imperialis, and stictica Engel for leonina as adopted in my
previous papers and this one. He agrees with me in the acceptance of parva
Schiner, which I have since sunk as a synonym of chrysame Walker, and in a
recent letter to me Dr. Engel has suggested the same synonymy, though he has
not had confirmation from Major Austen of the British Museum, to whom he has
written for information on the matter. Dr. Engel had seen only the type of
parva, which I had also seen, and he based his conclusions on the synonymy of
leonina and imperialis on the strength of an opinion expressed to him by Mr.
E. Seguy of the Paris Museum, where supposedly the Robineau-Desvoidy type
should be. We accept the determination of imperialis, but respectfully submit
that leonina, the type of which may be destroyed, judging from the Wiedemann
description, is quite distinct and is certainly identical with stictica Engel.

AMENIA CHRYSAME (Walker).
List Dipt. Brit. Mus., pt. 4, 1849, 866: Malloch, Proc. Linn. Soc. N.S.W., 55,
pt 2; 1980; 101.
Resembles a small Jeonina in general colour, with the same dark sides on
the upper half of back of head, but the upper postocular orbits are silvery-white,
not yellow, the hairs and bristles of the head are all black, the proclinate frontal,

76 NOTES ON AUSTRALIAN DIPTERA. XXxXiii,

the acrostichal, and anterior two pairs of the postsutural dorsocentral bristles are
much stronger, and there are submedian silvery spots on 2nd tergite.

In addition to the records I have already published, I have before me now a
specimen from Katoomba, Blue Mts., N.S.W., 3,400 feet (Dodd).

AMENIA DUBITALIS Malloch.
Proc. Linn. Soc. N.S.W., 52, pt. 3, 1927, 343.
I have seen no further material of this species.

AMENIA NIGROMACULATA Malloch.
Proc. Linn. Soc. N.S.W., 54, pt. 4, 1929, 286.
I have seen only the original male and female specimens of this species.

AMENIA SEXPUNCTATA, Nl. SD.

6, 9.—Black, with a violet-blue lustre, most pronounced on the posterior
portion of the thorax, and on the abdomen. Head orange-yellow, ocellar spot
fuscous, interfrontal stripe darker than the frontal orbits, the latter and the
parafacials and genae densely golden-yellow-dusted; antennae and palpi orange-
yellow; proboscis black; genal and lower occipital hairs yellow, the others and
bristles black; postocular orbits concolorous with genae. Thorax black, with a
distinct blue or violet-blue lustre, most pronounced posteriorly and on scutellum,
anterior margin of mesonotum with white dust, most distinct on two short stripes
just outside of the dorsocentral series and two more conspicuous stripes against
the humeri, the region between the dorsocentrals less densely dusted; postsutural
region with the usual two white-dusted spots, one supra-alar and the other
postalar, the central region hardly dusted; pleura with two silvery-white-dusted
spots, one on the mesopleura and the other on the sternopleura. Abdomen more
evenly violet-blue than the thorax, with a pair of white-dusted spots on disc of
each tergite from second visible one to fourth, largest on fourth, and with a small
spot of the same colour on the curve of second and another on fourth; all hairs
and bristles black. Legs black, the femora slightly blue-tinged. Wings hyaline,
infuscated, but not black, at bases. Calyptrae black, with the connecting part
white. Halteres brownish-black.

Frons of male about as wide in front of ocelli as third antennal segment, the
interfrontalia not entirely obliterated, the orbits with a series of fine inner
marginal bristles and laterad of these numerous microscopic hairs, the ocellars
minute and only the inner pair of verticals well developed; frons of female nearly
one-third of the head-width, the orbits narrowed behind, at middle each is
distinctly wider than the interfrontalia, and has one or two strong proclinate
outer bristles in addition to the same armature as the male; all four vertical
bristles well developed; facial carina broad, sometimes slightly sulcate centrally,
vibrissae situated at about half the length of third antennal segment above mouth
margin; parafacial about half as wide as height of gena, the latter about as
high as length of antenna; arista plumose. Presutural acrostichals moderately
strong, the anterior two pairs of postsutural dorsocentrals shorter than the
posterior pairs; scutellum slightly concave or emarginate on each side of the
apical pair of marginal bristles. Second visible tergite with a pair, the third
and fourth each with a complete series of apical bristles. Costa and the radial
veins not abnormally curved, third vein quite copiously setulose on basal third
of the section basad of the inner cross-vein. Length, 14-15 mm.

~]

-~]

BY J. R. MALLOCH.

Type, male, allotype, and two male paratypes, Palmerston, N.T., October,
1908 (Oldenberg coll., Deutsches Entomologisches Institut). One male paratype
will be sent to the Australian Museum through, the kindness of Dr. Walther
Horn.

I append below a key to the species of the genus known to me at this time.

Key to the Species.

1. Abdomen with the first visible tergite deep black, the others densely coated with
greyish-white dust through which a green tinge shows from the underlying
surface, second tergite with five deep black marks, a spot in centre, a small
one on each side at the lateral curve, and a small mark on each extreme lateral
edge below, third tergite with two additional black spots, each about as large
as the central one, situated between the latter and the small one at the lateral
curve, fourth tergite with a large central subtriangular black apical mark, its
Inibael, saMEHASAy JorOEV! GogcsopdouamdeoroodoboouDHoOoUnObGOS nigromaculata Malloch.

Abdomen either metallic-blue, blue-green, or violet-blue, with small isolated silvery-
white-dusted spots on some of the tergites, not with deep black spots on a
erevaish-wilte-dusteGdmenOun mere -alacrroksiotsctemstcbeion ei Mcrae olelolieNel ava: of (oll Fellailel «ale tala) 2

Species deep violet-blue or blue-black, the mesonotum with the usual three silvery
spots on each side and at least two white-dusted presutural vittae, and the
abdomen with a pair of silvery-white-dusted submedian spots on visible tergites
BiUno-ch VOWED coos cgonood accu bd Fosco comodo Oo.ooe oobOoua0> sexpunctata, n. sp.

Species more brightly coloured, generally metallic blue or green, and never with
a pair of submedian silvery-white-dusted spots on the third visible tergite of
GE LD CLOTIMETIG PY apes Seer on mehr tated ci ah cect iench serie Senrer ee aitatay sivel syieteeitey ahr staite Nalreltetioy site ve! sick es evs e:reivei’s 3

Mesonotum without evident submedian white-dusted presutural vittae; abdomen
without a trace of white submedian spots on the second visible tergite, only the
one on each lateral curve present; frons of male at narrowest point much more
than twice as wide as distance across posterior ocelli; presutural acrostichals
not noticeably differentiated from the adjoining hairs ..................... 4

Mesonotum with a pair of very evident white-dusted submedian presutural vittae 5

4. Ocellar bristles well developed and widely divergent in male, the frons at vertex in

same sex five or six times as wide as the distance across posterior ocelli ......

6.08’ 0 C.018 Gio elo SuOReAC RONG et CianonG I colen eReaniCOISL CR ce Sean Ae aioe imperialis Robineau-Desvoidy.
Ocellar bristles almost indistinguishable in the male, the frons at vertex in same
sex not four times as wide as the distance across posterior ocelli ..............
Seem Maye Mans ceiaiiel ore WATGNeS o's 2) ajcol att ahepote ed emarichs couch aiieheuenspawevenec crete wire rene dubitalis Malloch.

». Anterior postsutural dorsocentral bristles short and weak; postocular orbits very
little paler on upper half than below; second visible abdominal tergite with a
pair of submedian white-dusted spots, sometimes faint .... leonina (Fabricius).
Anterior postsutural dorsocentral bristles quite long and strong; postocular orbits
silvery-white on upper half, contrasting sharply with the yellow-dusted lower
portion; second visible tergite of the abdomen without submedian white-dusted
spots, only the usual one at each lateral curve .......... chrysame (Walker).

bo

(Je)

Genus PSEUDOTRICHOPODA, N.g.

This genus belongs to a tribe which is so far unknown from Australia. The
principal characters that distinguish the group consist of the bare parafacials
and arista, elongate narrow abdomen which has at least five tergites visible
when seen directly from above, the reduced tergal bristles, bare prosternum and
propleura, and the very much widened posterior portion of the lower calypter
which is almost transverse on its posterior margin and very noticeably emarginate
on its outer edge. The posterior margin of the thorax above the hind coxae and
below the base of the abdomen is rather high and more chitinized than usual
except in the Cylindromyiinae, but there are some fine transverse rugae present.
The Cylindromyiinae have, in addition to the more heavily chitinized posterior
margin of the thorax noted above, the lower calypter narrower and distinctly
rounded on hind margin, and without an emargination of the outer side. In

78 NOTES ON AUSTRALIAN DIPTERA. XXxXiii,

the structure of the lower calypter, and also in the more prominent knob just in

front of the wing on the pleura, the group strongly resembles the Phasiinae, but

the species are all much more. slender and the apical bristles on the abdominal

tergites, though weaker than in most Tachinidae, are usually distinguishable,

moderately strong in the present genus. The first posterior cell ends almost in

the apex of the wing and is closed in the margin, almost short stalked.
Genotype, the following species.

PSEUDOTRICHOPODA VARIPES, Nn. sp. Text-fig. 1.

$.—Head whitish-yellow, frons deep velvety brownish-black except on the
orbits, the latter becoming dark on the linear upper portions, back of head
infuscated above but densely whitish-grey-dusted so that the ground colour
appears pale, antennae brownish-black, apex of second segment reddish-yellow;
aristae dark brown; palpi pale orange-yellow; proboscis dark brown; frontal
hairs and bristles, the upper postocular cilia, and the hairs and bristles on
vibrissal angle black, the lower postocular cilia and the beard and genal hairs
whitish-yellow; frontal orbits and upper portions
of the parafacials yellowish-dusted, the face and
cheeks white-dusted. Thorax fuscous, the pleural
selerites more yellowish along the sutures, grey-
dusted, and with an oblique streak below wing base
and a paler mark on anterior half of mesopleura
darker than the remainder of surface; mesonotum
with dense yellowish-grey dust upon which there
are two deep black fasciae, one on anterior two-
thirds of presutural and the other on the same
extent of the postsutural area; scutellum deep black,
merging into rufous-yellow at apex; pleural hairs
whitish-yellow, remainder of hairs and bristles black.
Fig. 1—Head of Pseudotri- Abdomen black, first visible tergite translucent
chopoda varipes, from the ‘i 3

rt testaceous-yellow, except on a narrow dorsocentral
vitta, second tergite with a large spot of the same
pale colour on each side of anterior third, third tergite with two similar but much
smaller pale spots in front, the black portions slightly greyish-dusted, more
noticeably so apically. Legs orange-yellow, fore femora except their bases, the
extreme apices of mid and hind femora, and all of the tibiae and tarsi of all pairs
black. Wings very noticeably browned, especially on apical ccstal portion, the
veins golden-brown, and a slight tinge of same colour on the membrane on costal
half basally when seen from the tip and at a low angle. Calyptrae brownish-
yellow, the upper one white and the inner portion of the lower one paler yellow.
Halteres brownish-yellow.

Eyes bare; more than twice as high as long at centre; frons at vertex about
one-sixth of the head-width, twice as wide at anterior margin, the orbits linear
above and becoming gradually wider in front, with a series of fine incurved inner
marginal bristles on entire length which decrease in length above; ocellar bristles
quite well developed, longer and stronger than the inner verticals, proclinate and
slightly divergent, the inner verticals not longer than a pair of parallel bristles
situated behind the ocelli; parafacials with two or three fine black hairs opposite
the second antennal segment; head in profile as Figure 1. Thorax with 2+ 3 dorso-
centrals, only two bristles on the presutural lateral area, one postsutural intra-

BY J. R. MALLOOH. A9

alar, one very inconspicuous pair of short acrostichals just in front of the suture,
and the prealar very small and fine; sternopleurals 1 + 1; seutellum with four
marginal bristles, and the short hairs descending sides but not at apex; no hairs
below lower calypter. Abdomen slender, second tergite slightly longer than first
and third visible, the fifth shortest and in type depressed on dise but possibly
abnormal, surface hairs strong and decumbent, the apical central bristles distin-
guishable on all tergites but the first visible one, the fourth with an almost
complete series, fifth with but a few short setulae on dorsum; fifth sternite broadly
rounded at apex, not cleft, with a number of fine hairs and about six strong but
rather short bristles in a group on each side of median line at apex; hypopygium
retracted in type, the basal segment rounded and without hairs or bristles. Legs
rather strong, the hind tibiae thicker than usual in unrelated groups, but without
any scale-like anterodorsal bristles, the fore femur without distinct posteroventral
bristles, mid pair with one or two ventral bristles on basal half, the hind pair
with some similar bristles and but one bristle near apex on the anteroventral
surface; fore tibia with a short submedian posterior bristle; fore tarsi slender
at base, the apical two or three segments slightly widened; mid tibia with one
anterodorsal, one posterodorsal, and one strong ventral bristle, all near middle;
hind tibia with the short black hairs rather dense, and one anteroventral, and
two anterodorsal and posterodorsal bristles. Wings about three times as long
as wide, apices narrowly rounded, the first posterior cell ending in apex and
with a very short stalk, inner cross-vein slightly beyond middle of discal cell but
distinctly proximad of level of apex of first vein, third vein with one or two
hairs at base; bend of fourth vein subangular. Length, 10 mm.

Type, Cairns, N. Qld. (Collection Oldenberg). In the Deutsches Entomol-
gisches Institut, Berlin-Dahlem, Germany,

This genus differs from any other in the group in which I have placed it by
the lack of lanceolate anterodorsal bristles on the hind tibiae, the possession of
but two sternopleural bristles, the closed first posterior cell, and in having a few
hairs on the upper part of the parafacials.

As far as we know all the closely related genera are parasitic upon Hemiptera.

REVISION OF AUSTRALIAN LEPIDOPTERA. OECOPHORIDAE. II.
By A. JEFFERIS TURNER, M.D., F.E.S.
[Read 26th April, 1938.]

17. Gen. TELANEPSIA, 0.g.
Tyndaveyos, a distant cousin.

Head with loosely appressed scales. Tongue present. Maxillary palpi minute,
appressed to tongue. Labial palpi with second joint reaching base of antennae,
moderately thickened with loosely appressed scales; terminal joint shorter than
second, slender, acute. Antennae without basal pecten. Posterior tibiae with long
hairs on dorsum. Forewings with 2 from closely before angle, 7 and 8 stalked, 7
to costa. Hindwings broadly ovate; 3 and 4 connate, 5 from middle of cell.

Apparently allied to the New Zealand genus Gymnobathra, and if so an
interesting discovery. In the type specimen the left forewing is abnormal in
having 7 and 8 coincident, and 3 and 4 shortly stalked. Gymnobathra differs in
neuration by the origin of 2 of forewings from long before angle.

118. TELANEPSIA ORICALLA, N. SD.
éptxaddos, A Mountain beauty.

g. 22 mm. Head white. Palpi with second joint reaching base of antennae,
terminal joint three-fourths; white. Antennae fuscous. Thorax fuscous; apices
of tegulae and a large posterior spot white. Abdomen ferruginous-fuscous;
apices of segments grey-whitish. Legs fuscous, tibiae and tarsi with whitish
rings; posterior pair mostly whitish. Forewings moderate, not dilated, costa
gently arched, apex pointed, termen slightly rounded, strongly oblique; white; a
basal fuscous costal spot; a rather narrow dark fuscous fascia from two-fifths
costa to one-third dorsum, anterior edge straight, posterior sometimes irregular;
an irregular fascia from two-thirds costa to tornus, narrow on margins, strongly
dilated in disc with a short anterior truncate projection, brownish-fuscous; a
large triangular brownish-fuscous subapical spot; some dark fuscous dots around
apex and on termen; cilia whitish-ochreous. Hindwings whitish with a slight
grey suffusion towards apex and termen; cilia ochreous-whitish.

New South Wales: Barrington Tops in December; one specimen received from
Mr. G. M. Goldfinch, who has the type.

18. Gen. BAaryzANCLa, N.g.
Bapufayxdos, with heavy sickles.

Tongue present. Palpi very long, recurved; second joint three times length
of face, much thickened with appressed scales, somewhat rough anteriorly;
terminal joint about two-thirds, slender, acute. Antennae without basal pecten;
ciliations in male minute. Forewings with 2 and 8 closely approximated, connate,
or stalked, 7 to costa. Hindwings elongate-ovate; 3 and 4 connate, 5 from below
middle of cell.

BY A. J. TURNER. $1

A derivative of Leptocroca, differing in the palpi and antennae. Type,
B. dysclyta.

119. BARYZANCLA DYSCLYTA, N. Sp.
6vox\uTos, inglorious.

3, %. 18-20 mm. Head and thorax grey. Palpi grey mixed with fuscous;
posterior surface of terminal joint whitish. Antennae grey, becoming fuscous
towards apex. Abdomen grey; tuft grey-whitish. Legs fuscous. Forewings sub-
oblong, rather narrow, costa slightly arched to middle, thence straight, apex
round-pointed, termen oblique; grey; stigmata fuscous, minute or obsolete; first
discal at one-third, plical beyond it, second discal before two-thirds; cilia grey.
Hindwings and cilia pale grey.

Western Australia: Kalamunda, near Perth, in December; five specimens
received from Mr. W. B. Barnard, who has the type.

120. BArRyYZANCLA ITHYGRAMMA, Nl. SD.
iOvypaumos, With straight markings.

3, ¢. 18 mm. Head and thorax grey. Palpi grey; anterior edge of second
joint, and terminal joint towards apex, fuscous. Antennae fuscous. Abdomen
ochreous-whitish, towards apex grey; tuft ochreous-whitish. Legs grey; posterior
pair ochreous-whitish. Forewings narrow, suboval, costa moderately arched, apex
pointed, termen very obliquely rounded; grey-whitish irrorated with fuscous;
markings dark fuscous; a narrow streak along fold to one-third; a discal dot at
one-third, another at two-thirds, between them a short longitudinal streak; some
dark fuscous irroration on veins towards termen; cilia pale ‘grey with some
dark fuscous points. Hindwings and cilia pale grey.

Western Australia: Mt. Dale in January; two specimens received from Mr.
W. B. Barnard, who has the type.

19. Gen. Hopnostrea Meyr.
Exot. Micro., i, p. 235.

Palpi rather short, curved, ascending; second joint not nearly reaching base
of antennae, thickened with appressed scales, terminal joint nearly as long as
second, rather stout. Antennae in male stout, shortly ciliated, basal joint concave
beneath, forming a small eyecap, pecten present. Forewings with 2. and 3 stalked,
7 to costa. Hindwings elongate-ovate; neuration normal.

Monospecific, allied to Leptocroca, differing in the peculiar antennae.

121, ochroma Meyr., P.L.S.N.S.W., p. 781 (Brisbane; Sydney; Beaconsfield).

20. Gen. GoNIOBELA, 0.g.
ywrvioBedos, With angled palpi.

Tongue present. Palpi moderately long, curved, ascending; second joint not
reaching base of antennae, thickened with appressed scales and dilated at apex
into a small anterior apical tuft; terminal joint shorter than second. Antennae
with basal pecten; ciliations of male rather long. Forewings narrow, elongate;
2 and 3 stalked, 7 to costa. Hindwings elongate-ovate; 3 and 4 connate, 5 from
middle of cell,

Type, G. astatopis. A derivative of Leptocroca, from which it may be readily
distinguished by the palpi.

Three species: 122, astatopis, n. sp. (Macpherson Range).—123, nonymopis,
n. sp. (Toowoomba) .—124, idiospila, n. sp. (Mt. Gregson, N.S.W.).

i

82 REVISION OF AUSTRALIAN LEPIDOPTERA. OECOPHORIDAE. ii,

122. GoNIOBELA ASTATOPIS, 0. SD.
aoTratwris, variable.

g. 19-21 mm. Head and thorax fuscous-brown. Palpi brown. Antennae
brown, basal joint fuscous; ciliations in male 3. Abdomen grey. Legs fuscous;
posterior pair whitish-ochreous. Forewings narrow, elongate-oval, costa rather
strongly arched, apex rounded, termen very obliquely rounded; fuscous-brown; a
sub-basal median fuscous suffusion; discal dots approximated at about two-fifths
and three-fifths, plical before first discal; a curved subterminal line of fuscous
dots commencing beneath three-fourths costa; an obscure series of costal dots
from middle to apex; cilia fuscous-brown, bases barred with fuscous. Hindwings
and cilia whitish, faintly ochreous-tinged.

This species is variable. In a second example the antennae, head, thorax,
and basal third of forewings except dorsum are brown-whitish. In a third the
forewings are pale fuscous without markings.

Queensland: National Park (3,000 ft.) in March; three specimens.

123. GONIOBELA NONYMOPIS, Nl. SD.

vwvunwres, insignificant.

do. 17 mm. Head and palpi whitish. Antennae whitish; ciliations in male 3.
Thorax whitish. Abdomen grey; tuft whitish. Forewings narrow, elongate-oval,
costa rather strongly arched, apex round-pointed, termen very obliquely rounded;
whitish with slight fuscous suffusion near dorsum and tornus; cilia whitish.
Hindwings and cilia whitish.

Queensland: Toowoomba in April; one specimen.

124. GONIOBEILA IDIOSPILA, N. Sp.
l6cogmcAos, With a peculiar spot.

6. 18-20 mm. Head and thorax grey. Palpi fuscous; inner surface of second
joint whitish. Antennae grey; ciliations in male 3. Abdomen grey; tuft ochreous-
whitish. Legs fuscous; posterior pair ochreous-whitish. Forewings elongate-oval,
costa moderately arched, apex pointed, termen very obliquely rounded; grey with
fuscous markings; a large triangular spot on costa at one-third; a sub-basal
median dot; a white spot above tornus surrounded by a considerable fuscous
suffusion; a transverse discal mark at three-fifths touching this; a series of dots
from beneath midcosta, sharply bent before apex and continued parallel to termen;
cilia grey-whitish with a median series of fuscous bars. Hindwings and cilia
whitish-grey.

Variable; in a second example head, thorax, and basal third of forewings
except dorsum are whitish.

New South Wales: Mt. Gregson in March; two specimens received from. Mr.
G. M. Goldfinch, who has the type.

21. Gen. Lreprocroca Meyr.
Proc. Linn. Soc. N.S.W., 1885, p. 775. Type, L. sanguinolenta Meyr.

Palpi moderately or very long, curved, ascending; second joint reaching or
exceeding base of antennae, thickened with appressed or loosely appressed scales;
terminal joint shorter or rarely as long as second (three-fifths to one). Antennae
with basal pecten; ciliations in male short or long (1 to 5). Forewings with
2 and 3 connate or stalked, 7 to costa. Hindwings elongate-ovate; 3 and 4
connate, 5 from below middle of cell.

BY A. J. TURNER. 83

An endemic derivative of Borkhausenia, easily distinguished when 2 and 3
of forewings are stalked, less easily when these veins are almost connate. I
include here the two species referred to Pawronota Low., though with some
doubt, as I have not seen either of them. I include also Guestia uniformis Meyr.
It is true that in some examples of this species 2, 3 and 4 of forewings are stalked,
but in others 4 arises from the cell. I have examined a specimen of tetralychna
Low., the type of Ardozyga Low., and find that it is a Protolechia. ‘That genus
therefore disappears.

Forty species: 125, synaptospila, n. sp. (Mt. Tambourine) .—126, stenophanes,
n. sp. (Beaconsfield, Macedon, Strahan).—127, niphadia, Meyr., P.L.S.N.S.W., 1885,
p. 795 (Mackay to Sydney) .—128, eucentra, Turn., Proc. Roy. Soc. Tas., 1927, p. 139
(Tasmanian Mts.).—129, notospila, n. sp. (Sydney).—130, sanguwinolenta, Meyr.,
P.L.S.N.S.W., 1885, p. 775 (Brisbane to Melbourne; Adelaide).—131, comarcha
Meyr., Hxot. Micro., ii, p. 367 (Pinnaroo, §.A.).—?+132, epimicta Meyr., P.L.S.N.S.W.,
1885, p. 786 (Deloraine, Tas.).—133, platynephela, n. sp. (Toowoomba) .—134,
chersomicta Meyr., Exot. Micro., ii, p. 308 (Brisbane, Mt. Tambourine).—135,
polioleuca, n. sp. (Cape York).—136, todes Low., Tr.R.S.S.Aust., 1901, p. 94
(Adelaide, Mt. Lofty) .—137, ischnota Low., ibid., 1903, p. 226 (Broken: Hill).—
138, actinipha Low., ibid., 1901, p. 95 (Broken Hill).—7139, grammocentra Meyv..
Exot. Micro., ii, p. 367 (Duaringa).—140, delosticha Low., Tr.R.S.S.Aust., 1915,
p. 483 (Broken Hill).—141, megaloplaca Low., P.L.S.N.S.W., 1900, p. 46 (Broken
Hill) —142, symmadelpha Low., Tr.R.S.S.Aust., 1915, p. 483 (Broken Hill).—
143, thermoloma Low., ibid., 1901, p. 96 (Broken Hill).—144, lasioprepes Low.,
P.L.S.N.S.W., 1915, p. 484 (Broken Hill).—145, adelphodes Low., Tr.R.S.S.Aust.,
1893, p. 178 (Gisborne, Adelaide). —146, spanioleuca, n. sp. (W.A.: Denmark) .—
147, athletis Meyr., P.L.S.N.S.W., 1887, p. 961 (Mt. Lofty).—148, sphaleropis Meyr.,
Tr.R.S.S.Aust., 1902, p. 168 (Beaconsfield, Gisbornie).—149, adoxodes, n. sp.
(Macpherson Range).—150, clepsiphanes, n. sp. (W.A.: Mundaring).—151,
dryinodes Meyr., P.L.S.N.S.W., 1888, p. 1565 (Melbourne, Adelaide) = peladelpha
Low., Tr.R.S.S.Aust., 1894, p. 101 (W.A.: Kalgoorlie).—152, eurybapta Low.,
ibid., 1908, p. 117 (Broken Hill).—153, zophosema, Low., ibid., 1905, p. 108 (Broken
Hill). —154, amydrosema, Low., ibid., 1903, p. 227 (Mt. Wellington) .—7155, nicaea
Meyr., ibid., 1902, p. 147 (Tasmania).—156, dysopta, n. sp. (Macpherson Range) .—
157, meselectra, Meyr., ibid. 1902, p. 148 (Duaringa, Gympie, Brisbane,
Toowoomba).—7158, ophthalmias Meyr., P.L.S.N.S.W., 1887, p. 950 (W.A.:
Albany). —7159, pseudopis Meyr., Exot. Micro., ii, p. 368 (W.A.: Busselton) —
160, eusema Low., P.L.S.N.S.W., 1900, p. 418 (Broken Hill).—161, balia, n. sp.
(Bunya Mts.).—162, chaetophora, n. sp. (Tweed Heads).—163, wniformis Meyr.,
ibid., 1885, p. 781 (Bathurst, Mt. Canoblas, Mt. Lofty)—164, caenosa, n. sp.
(Bourke).

125. LEPTOCROCA SYNAPTOSPILA, 1. SP.
ouvatToortdos, With conjoined spots.

6- 15-16 mm. Head pale ochreous. Palpi with second joint not reaching
base of antennae, terminal joint three-fifths; fuscous mixed with pale ochreous.
Antennae grey; ciliations in male nearly 1. Thorax fuscous. Abdomen whitish-
ochreous. Legs fuscous; posterior pair whitish-ochreous. Forewings narrow,
oval, costa rather strongly arched, apex acute, termen extremely oblique; whitish-
ochreous with fuscous irroration and markings; an ill-defined fuscous blotch on
base of costa; first discal at one-third, plical slightly beyond it, both enlarged

84 REVISION OF AUSTRALIAN LEPIDOPTERA. OECOPHORIDAE. ii, ©

and confluent forming a slightly oblique transverse bar; second discal very
obscure at two-thirds, a more distinct spot before and beneath it; sometimes
cbscure suffused costal spots at three-fourths and near apex; cilia ochreous-
whitish. Hindwings elongate-ovate; pale grey; cilia pale grey.

Queensland: Mt. Tambourine in November; two specimens.

126. LeEPpTocROcCA STENOPHANES, 0. SD.
oTevoparns, Narrow.

3, . 16-18 mm. Head brown-whitish. Palpi with second joint just reaching
base of antennae, terminal joint three-fifths; ochreous-whitish; ciliations in male 1.
Thorax whitish-grey. Abdomen in male whitish, in female grey; tuft ochreous-
whitish. Legs grey; posterior pair ochreous-whitish. Forewings narrow, elongate,
oval; costa slightly arched, apex pointed, termen very obliquely rounded; whitish
more or less densely irrorated with pale fuscous; an outwardly curved fuscous
line from one-third costa to one-third dorsum, sometimes very distinct; a fuscous
discal dot at two-thirds; a series of minute fuscous dots very near to termen and
apical one-fourth of costa; cilia whitish with fuscous irroration. Hindwings
elongate-ovate; in male whitish-grey, in female grey; cilia whitish.

Victoria: Beaconsfield and Macedon in November (Lyell). Tasmania: Strahan
in February (¢ type). Three specimens.

129. LerPTOCROCA NOTOSPILA, 1. Sp.
vwroomtAos, With dorsal spot.

3, @. 18-20 mm. Head and thorax pale grey. Palpi with second joint not
reaching base of antennae, terminal joint three-fifths; grey. Antennae grey;
ciliations in male 13. Abdomen and legs grey. Forewings narrow, costa strongly
arched, apex pointed, termen extremely oblique; 2 and 3 connate; pale grey
with scattered fuscous irroration mostly in distal two-thirds; an irregular sharply
defined dark fuscous blotch on dorsum from one-third to two-thirds, its apex
formed by plical stigma; beyond this a triangular spot on costa at one-third, its
apex formed by first discal; second discal at two-thirds, sometimes confluent
with a dot near termen above tornus; a series of minute fuscous dots close to
termen and apical half of costa; cilia grey. Hindwings elongate-ovate; pale
grey; cilia pale grey.

Very similar to L. eucentra Turn., but the head is grey, not whitish, and the
forewing lacks the basal and discal dots of that species.

New South Wales: Sydney (St. Mary’s) in September; two specimens
received from Mr. G. M. Goldfinch, who has the type.

133. LePprocRocA PLATYNEPHELA, N. Sp.
mhatuvepedos, broadly clouded.

9. 14-16 mm. Head ochreous-whitish. Palpi with second joint not reaching
base of antennae, terminal joint three-fifths; dark fuscous, second joint with post-
median and apical, terminal joint with median whitish rings. Antennae blackish
finely annulated with whitish. Thorax dark fuscous; tegulae and a posterior dot
whitish. Abdomen ochreous-grey. Legs whitish with dark fuscous irroration.
Forewings rather narrow, costa slightly arched, apex pointed, termen obliquely
rounded; 2 and 3 stalked; white rather densely irrorated with dark fuscous
so as to appear grey; but central area of disc suffusedly whitish; stigmata
fuscous, often indistinct, first discal at one-third, plical slightly beyond it, second
discal at two-thirds, a dark fuscous costal dot at middle and another beyond it:

BY A. J. TURNER. 85

a series of dark fuscous dots forming a line from three-fourths costa to tornus,
curved close to termen; cilia ochreous-whitish, bases irrorated with dark fuscous.
Hindwings and cilia grey.
The annulated palpi should be useful in distinguishing this obscure species.
Queensland: Toowoomba in October; five specimens received from Mr. W. B
Barnard, who has the type.

135. LEPTOCROCA POLIOLEUCA, 0. Sp.

moNwodeukos, greyish-white.

fg, 9. 14-15 mm. Head and thorax greyish-white. Palpi with second joint
exceeding base of antennae, terminal joint two-thirds; white, second joint with
subapical, terminal joint with basal and subapical blackish rings. Antennae
whitish; ciliations in male less than 1. Abdomen whitish. Legs grey-whitish.
Forewings suboblong, costa gently arched, apex rounded, termen very obliquely
rounded; greyish-white with markings and some scattered scales blackish; first
discal at one-third, plical well beyond it, second discal at two-thirds; a series
of dots close to margin along apical third of costa and whole of termen; cilia
white. Hindwings elongate-ovate; pale grey; cilia pale grey.

North Queensland: Cape York in October; five specimens received from Mr.
W. B. Barnard, who has the type.

136. Lerprocroca 1opEs Low.

I have examined a specimen so named from Coll. Lower. In this the
pecten is denuded, but a few scales are left, and 2 and 3 of forewings are
stalked. It is not the type, and there is therefore a possibility of erroneous
identification, but that iodes is referable to Schiffermuelleria appears to me very
improbable.

146. LeprocrocaA SPANIOLEUCA, 0. SD.
omaviovevxos, Scantily white.

3, 9. 20-22 mm. Head and thorax grey. Palpi with second joint exceeding
pase of antennae; fuscous with a more or less pronounced whitish subapical band,
inner surface mostly whitish; terminal joint three-fourths, fuscous. Antennae
grey; ciliations in male 1. Abdomen grey; tuft grey-whitish. Legs fuscous;
posterior pair except tarsi whitish. Forewings suboblong, costa gently arched,
apex rounded, termen obliquely rounded; whitish with fine fuscous irroration,
appearing grey; a short fuscous streak from base of costa; a short longitudinal
dark fuscous streak before middle of disc, sometimes obsolete, succeeded by a
whitish dot, and this by a dark fuscous discal dot beyond middle; an outwardly
curved fuscous line from three-fourths costa to tornus, sometimes indistinct; beyond
this a series of short radiating streaks to termen; cilia grey, apices whitish.
Hindwings and cilia pale grey.

Western Australia: Denmark in March and April; eight specimens received
from Mr. W. B. Barnard, who has the type.

149. LerPTrocROocA ADOXODES, N. Sp.
“ad0éwdys, Obscure.

g. 18 mm. Head and thorax fuscous. Palpi with second joint not reaching
base of antennae, terminal joint two-thirds; fuscous, apices of second and terminal
joints ochreous-whitish. Antennae dark fuscous with fine whitish annulations;
Ciliations in male 2. Abdomen reddish-brown; apices of terminal segments and

86 REVISION OF AUSTRALIAN LEPIDOPTERA. OECOPHORIDAE. 1i,

tuft grey. Legs fuscous with ochreous-whitish rings; posterior pair mostly
ochreous-whitish. Forewings narrow, oblong, costa nearly straight, except at base,
apex rectangular, termen slightly rounded, slightly oblique; whitish with dense
fuscous irroration, which is uniform except over centre of disc and before apex,
which are paler; stigmata dark fuscous, obscure, first discal at one-third, plical
beneath it, second discal at two-thirds; a dark streak connects second discal with
apex, and a large round spot with tornus, both are ill-defined; cilia pale grey
with two rows of fuscous points. Hindwings and cilia whitish-grey.

Queensland: Springbrook, Macpherson Range, in October; one specimen
received from Mr. W. B. Barnard.

150. LEPTOCROCA CLEPSIPHANES, Nl. Sp.
xvevidarns, of misleading appearance.

9. 20 mm. Head, thorax, and abdomen grey. Palpi with second joint
exceeding base of antennae, terminal joint two-thirds; pale grey with some
fuscous irroration. Antennae grey. Legs grey; posterior pair whitish. Fore-
wings elongate, slightly dilated posteriorly, costa rather strongly arched, apex
round-pointed, termen obliquely rounded; pale grey with fuscous irroration;
markings dark fuscous; first discal forming a short narrow transverse discal
mark at one-third, plical beneath it, minute, second discal before two-thirds, a
short longitudinal streak between’ and above discals; veins in terminal area
partly outlined with dark fuscous; cilia grey-whitish with some fuscous points.
Hindwings broadly ovate; pale grey; cilia pale grey.

An obscure form, which looks like a Hulechria. Apparently a winter species.
Nearest L. sphaleropis.

Western Australia: Mundaring in June; one specimen received from Mr.
J. Clark.

156. LerprocRocA DYSOPTA, N. sp.
duvcomtos, obscure.

?. 18 mm. Head and thorax brown, with fuscous irroration. Palpi with
second joint exceeding base of antennae, terminal joint five-sixths; brown with
fuscous irroration. Antennae brown with fuscous annulations. Abdomen brown.
Legs fuscous with whitish-ochreous rings on tibiae and tarsi; posterior pair mostly
whitish-ochreous. Forewings elongate, costa slightly arched, apex rounded, termen
obliquely rounded; 2 and 3 connate or nearly so; brown sparsely irrorated with
fuscous; stigmata fuscous, first discal at one-fourth, followed by one or two
dots in a line, second discal beyond middle, plical slightly beyond first discal;
cilia brown. Hindwings elongate-ovate; pale grey; cilia pale grey.

This obscure species should be recognizable by its long palpi and general
brown coloration.

Queensland: National Park (3,000 ft.) in November; one specimen.

161. LeprocRocaA BALIA, Nn. Sp.
Badwos, spotted.

9. 22 mm. Head and thorax whitish-grey. Palpi with second joint exceeding
base of antennae, terminal joint one-half; whitish, second joint with base of
external surface and some scattered scales fuscous, terminal joint fuscous
anteriorly. Antennae fuscous becoming grey towards base, but basal joint fuscous.
Abdomen grey, apices of segments grey-whitish. Legs whitish; anterior pair
fuscous, tibiae and tarsi with whitish rings. Forewings suboval, costa strongly

BY A. J. TURNER. 87

arched, apex rounded, termen obliquely rounded; whitish-grey with some grey
suffusion and blackish markings; a streak from base along fold ending in an
elongate spot at one-fifth; discals small but distinct, first at one-third, second at
about middle; costal dots at one-fifth and three-fifths, and three before apex;
a fine dotted line from two-thirds costa, outwardly oblique, rather sharply angled
in middle, ending on dorsum before tornus; some terminal dots; cilia whitish-
grey. Hindwings grey; cilia grey-whitish.

Queensland: Bunya Mts. (3,000 ft.) in May; one specimen received from
Mr. W. B. Barnard.

162. LEprocrocA CHAETOPHORA, Nl. SDP.
xXavropopos, hair-plumed.

dg. 22-24 mm. Head and thorax fuscous. Palpi with second joint exceeding
base of antennae, terminal joint one-half; fuscous; extreme apices of second
and terminal joints whitish. Antennae fuscous; ciliations in male 1. Abdomen
fuscous; apices of segments and tuft ochreous-whitish. Legs fuscous; tibiae and
tarsi with ochreous rings. Forewings moderately broad, dilated posteriorly, costa
rather strongly arched, apex rounded, termen obliquely rounded; 3 and 4 out
of 2; whitish densely and uniformly irrorated with fuscous; markings obscure,
dark fuscous; discals approximated, first beyond one-third, second before two-
thirds, plical before first; a longitudinal streak between base and first discal,
sometimes a pale spot immediately following second discal; a subterminal series
ot dots from two-thirds costa to before tornus; cilia whitish, bases with fuscous
irroration, a grey subapical line. Hindwings ovate; in male with a spreading
tuft of long hairs from base of dorsum; ochreous-whitish becoming grey-whitish
towards apex; cilia whitish with two faint grey lines.

This species combines the peculiar neuration found sometimes, but not
constantly, in L. (Guestia) uniformis, with the expanding hairtuft on hindwings
found in B. (Disselia) aleurota.

Queensland: Southport in June and July; two specimens received from Mr.
W. B. Barnard, who has the type.

164. LerprocrocA CAENOSA, N. Sp.
caenosus, muddy.

o. 22 mm. Head pale brown. Palpi with second joint just reaching base of
antennae, terminal joint three-fourths; pale brown, base of second joint and
anterior surface of terminal joint fuscous. Antennae fuscous-brown; ciliations
in male 1. Thorax fuscous-brown. Abdomen grey. Legs ochreous-whitish;
anterior pair fuscous. Forewings suboblong, costa gently arched, apex rounded,
termen obliquely rounded; fuscous-brown; stigmata pale reddish-brown, partly
outlined with fuscous, first discal at one-third, plical well before it, second discal
before two-thirds, larger; cilia fuscous-brown. Hindwings elongate-ovate; grey;
cilia grey.

New South Wales: Bourke (Helms Coll.); one specimen.

22. Gen, PHANEROLOPHA, D.g.
pavepodogos, with distinct crest.
Palpi long, ascending, recurved; second joint reaching base of antennae,
thickened with appressed scales, somewhat more thickened and rough towards
apex; terminal joint as long as second, slender. Antennae with basal pecten;

58 REVISION OF AUSTRALIAN LEPIDOPTERA. OECOPHORIDAF. ii,

in male shortly ciliated. Thorax with a posterior crest. Forewings with 2 well
separate, 7 to costa. Hindwings with 3 and 4 stalked, 5 from below middle of cell.

A development of Borkhausenia. New species may be expected from inland
districts.

165. PHANEROLOPHA PELAEOBAPHES, MN. SD.

patoBagpys, dusky-suffused.

fg. 18 mm. Head and thorax whitish-grey with fuscous irroration. Palpi
fuscous, inner surface of second joint whitish. Antennae fuscous; ciliations in
male 1. Abdomen grey; tuft whitish-ochreous. Legs fuscous; middle tibiae and
tarsi with whitish rings; posterior pair ochreous-whitish. Forewings elongate-
oval, costa strongly arched, apex round-pointed, termen very oblique; whitish-
grey; basal, dorsal, and subapical areas broadly suffused with fuscous, leaving a
pale spot above one-fourth dorsum, and an ill-defined pale median streak; first
discal beyond one-third or obsolete, second discal at two-thirds; sometimes obscure
fuscous streaks on veins in terminal area; cilia fuscous, apices whitish-grey.
Hindwings elongate-oval; grey, paler towards base; cilia grey.

Queensland: Stanthorpe in February; Mitchell in September; two specimens.
Type in Coll. Barnard.

23. Gen. TRINACONEURA, Ng.
Tpivaxoveupos, trident-nerved.

Palpi rather short, curved, ascending; second joint not nearly reaching base
of antennae, with appressed scales; terminal joint two-thirds. Antennae
with basal pecten; ciliations in male long. Forewings narrow; 2 and 3 separate,
7 to costa. Hindwings narrow, elongate-ovate; 3 and 4 stalked or rarely coincident,
5 nearly approximated or connate from lower angle of cell.

A derivative of Borkhausenia.

166. TRINACONEURA HOMOGYPSA, DN. SD.

ouoyuyos, uniformly whitish.

dS. 16 mm. Head and thorax whitish. Palpi whitish; apex of second joint
fuscous. Antennae whitish; ciliations in male 3. Abdomen pale ochreous-brown,
apices of segments, tuft, and underside whitish. Legs whitish; anterior pair
fuscous. Forewings narrow, elongate-oval, costa slightly arched, apex pointed,
termen very oblique; whitish with some scanty grey irroration in terminal area;
cilia whitish. Hindwings elongate-ovate; whitish; cilia whitish.

Queensland: Stradbroke Island in August; one specimen,

24. Gen. BoRKHAUSENTA.
Hb., Verz., p. 420; Meyr., Gen. Insect. Oecoph., p. 37. Type, B. minutella Linn.
from Europe.

Palpi rather short or moderately long, curved, ascending; second joint not
reaching or exceeding base of antennae; with appressed scales; terminal joint
usually shorter than second. Antennae with basal pecten; ciliations in male short
or long (3% to 5). Forewings with 2 separate. 7 to costa. Hindwings elongate-ovate
or ovate-lanceolate (sometimes almost lanceolate); 3 and 4 connate, or rarely
stalked or even coincident, 5 rarely from middle of cell, usually from below
middle.

A large and primitive genus. Meyrick records 37 Palaearctic species, 7
Nearctic, 4 Neotropical, but none from the Oriental region. They are numerous
in New Zealand and Eastern Australia, and, being mostly small and inconspicuous,

BY A. J. TURNER. 89

many more will be discovered. Meyrick infers that the genus developed in
Central Asia, while separated from the Indian peninsula, and spread through
North America and the Andes to Antarctica, and thence to New Zealand and
Australia.

Though vein 7 of forewings runs normally to the costa, in an occasional
specimen it may run to the apex on one or both sides. Consequently the genus
may be confused with Hulechria, if only a single example is examined. Of four
examples of B. semiota (the type of Crossophora), in two 3 and 4 of hindwings
were found connate, in two coincident. In two of B. gypsomicta, not nearly
related to semiota specifically, these two veins were coincident in both. Meyrick
notes similar variations in the hindwings of WMulechria. The species of
Borkhausenia fall into two natural groups, those with narrow forewings and ovate-
lanceolate hindwings, and those with broader forewings and elongate-ovate hind-
wings, but intermediate forms occur, and it would be a mistake to divide the
genus.

Highty-one Species: 167, leptophylla, n. sp. (Hungella).—168, chryseres Turn.,
Tr.R.S.S.Aust., 1898, p. 207 (Brisbane to Melbourne). = amphixantha, Low.,
ibid., 1904, p. 169; saltuosa Meyr., Exot. Micro., i, p. 172.—169, pentochra Low.,
Tr.R.S.S.Aust., 1894, pD. 102 (Sea Lake, HEucla).—170, tetratherma Low., ibid.,
1896, 165 (Glen Innes to Melbourne) .—171, semiota Meyr., P.L.S.N.S.W., 1885, 797
(Duaringa, Brisbane, Sydney) .—7172, taractis Meyr., Exot. Micro., i, 297 (Sydney).
—1173, lychnosema Meyr., P.L.S.N.S.W., 1885, p. 787 (Sydney; St. Helen’s, Tas.).—
174, erythrocephala Low., Tr.R.S.S.Aust., 1904, p. 169 (Broken Hill).—175, nyctora
Meyr., Hxot. Micro., i, p. 297 (Katoomba, Gisborne) .—176, homopela, n. sp. (W.A.:
Denmark) .—177, diaxesta Meyr., Arkiv f. Zool., xiv (15), p. 6 (Atherton).—178,
maculifera Low., P.L.S.N.S.W., 1899, p. 11 (Milmerran, Broken Hill).—7179,
oenopa Meyr., ibid., 1885, p. 796 (Quorn, S.A.).—180, scotiodes Meyr., Tr.R.S.S.Aust.,
1902, p. 151 (Adelaide).—181, poliocrana Meyr., P.L.S.N.S.W., 1885, p. 787
(Katoomba).—182, tanytricha, n. sp. (Macpherson Range).—183, phanerosticta,
n. sp. (Macpherson Range).—184, perigrapta, n. sp. (W.A.: Denmark) .—185,
chalcoteucta, u. sp. (Gisborne).—186, hypochaicha Meyr., ibid., 1885, p. 782
(Sydney, Gisborne, Hast Tasmania) .—187, oxypeuces Turn., Proc. Roy. Soc. Tas.,
1926, p. 141 (Beaconsfield, Vic., Bothwell, Tas.).—188, gypsopleura Turn.,
P.L.S.N.S.W., 1916, p. 338 (W.A.: Cunderdin) .—189, crymorrhoa Meyr., ibid., 1888,
p. 1669 (Sydney, Pt. Lincoln, Tasmania).—190, dolosella Wl1k., xxviii, p. 539;
Meyr., ibid., 1882, p. 539; = petrophanes Meyr., Exot. Micro., i, p. 162 (Tenterfield,
Neweastle to Gisborne).—191, desiccata Meyr., Exot. Micro., i, p. 296 (W.A.:
Waroona, Geraldton) .—192, gypsomicta, n. sp. (Stanthorpe).—7193, catochopis
Meyr., ibid., ii, p. 307 (Brisbane).—7194, asparta Meyr., T7.R.S.S.Aust., 1906,
p. 36 (Sydney, W.A.: Albany).—19$5, lechriomochla, n. sp. (Barrington Tops).—
7196, spodostrota Meyr., ibid., 1902, p. 173 (Katoomba).—197, nigripuncta, n. sp.
(Cradle Mt.).—7198, lagara Meyr., P.L.S.N.S.W., 1885, p. 783 (Rosewood, Sydney) .—
7199, reprobata Meyr., Exot. Micro., ii, p. 307 (Brisbane).—200, albipectinata,
h. sp. (Brisbane, Toowoomba) .—201, eremaea Meyr., P.L.S.N.S.W., 1885, p. 783
(Brisbane, Toowoomba, Glen Innes, Tasmania).—202, nubifera Meyr., ibid., 1885,
784 (Duaringa, Brisbane, Glen Innes, Ebor). = cyclozona Low., Tr.R.S.S.Aust., 1905,
p. 109.—203, serrulifera, n. sp. (Brisbane).—204, aleuwrota Meyr., P.L.S.N.S.W..
1885, p. 799 (Toowoomba to Mt. Lofty; Tasmania).—205, iulophylla, n. sp. (W.A.:
Albany) .—206, lymphatica, Meyr., ibid., 1885, p. 785 (Mt. Wilson to Mt. Lofty;

90 REVISION OF AUSTRALIAN LEPIDOPTERA. OECOPHORIDAE. ii,

Tasmania).—207, macroptera Turn., ibid., 1916, p. 338 (Mt. Kosciusko).—208,
gypsodes, n. sp. (W.A.: Perth).—209, tholopa Turn., ibid., 1916, p. 336 (Mt.
Tambourine, Macpherson Range).—210, misella, n. sp. (Macpherson Range).—
211, trivialis Meyr., Exot. Micro., i, p. 172 (Mt. Wilson, Gembrook, Beacons-
field, Tasmania).—212, vernilis, n. sp. (W.A.: Denmark).—213, cnecocrana,
n. sp. (Sydney).—214, flavipuncta, n. sp. (Cairns).—215, paurophylla Turn.,
P.L.S.N.S.W., 1916, p. 337 (Brisbane, Stradbroke I., Tweed Heads) .—216, lissoptera,
n. sp. (Cairns, Stradbroke I.).—217, leptocneca, n. sp. (Hungella).—218, asyneta
Meyr., ibid.¢ 1885, p. 795 (Brisbane to Sydney).—7219, liacta Meyr., Exot. Micro.,
ii, p. 367 (Brisbane).—220, thetias Meyr., P.L.S.N.S.W., 1885, p. 796 (Sydney, Mt.
Lofty, W.A.: Albany).—221, nephelella Turn., Tr.R.S.S.Aust., 1898, p. 212
(Nambour to Lismore) .—222, dichroa Low., ibid., 1893, p. 179 (Melbourne, Birchip,
Adelaide). = callioptis Low., ibid., 1903, p. 226.—223, zophodes Meyr., P.L.S.N.S.W.,
1885, p. 784 (Katoomba) .—224, basileuca, n. sp. (W.A.: Mt. Dale).—225, lechrio-
gramma, n. sp. (W.A.: Busselton).—226, anthemodes Meyr., ibid., 1885, p. 780
(Mt. Wilson to Beaconsfield). = tetraphaea Turn., ibid., 1916, p. 337 (Tasmania) .—
227, hemisphaerica Meyr., ibid., 1885, p. 780 (Cape York to Tweed Heads) .—
228, sphaeroides Turn., Tr.R.S.S.Aust., 1896, p. 31 (Brisbane).—7229, chroma-
tarcha Meyr., Exot. Micro., i, p. 232 (Sydney) —230, canephora Meyr., P.L.S.N.S.W..,
1883, p. 339 (Mt. Gambier; Tasmania).—231, sufurea Meyr., ibid., 1885, p. 786
(Sydney to Pt. Lincoln; W.A.: Albany).—7232, hilaropa Meyr., ibid., 1888, p. 1672
(W.A.: Perth, York).—233, protadelpha Meyr., ibid., 1888, p. 1672 (Sea Lake;
W.A.: Perth, Busselton, Cunderdin) —234, cosmanthes Meyr., ibid., 1888, p. 1671
(W.A.: Geraldton ).—235, mesozona Low., Tr.R.S.S.Aust., 1908, p. 225 (Stawell). —
236, eurrhoa Meyr., P.L.S.N.S.W., 1885, p. 789 (Toowoomba to Gisborne; Mt.
Lofty: Tasmania).—7237, phthorodoxa Meyr., ibid., 1885, p. 794 (Sydney,
Katoomba) .—238, psaritis, n. sp. (Macpherson Range).—239, achroa Meyr., Exot.
Micro., i, p. 232 (Adelaide).—240, xuthochroa, n. sp. (Hobart).—241, pelophanes,
n. sp. (Tasmania).—242, acalles Turn., Proc. Roy. Soc. Tas., 1926, p. 139 (Mt.
Wellington). = silicolor Turn., ibid., 1926, p. 140.—+243, aetodes Meyr., P.L.S.N.S.W.,
1888, p. 1673 (Mt. Lofty) —244, centrosticha, n. sp. (Gisborne) .—245, nephotypa,
n. sp. (Katoomba) .—246, trichoceros, n. sp. (Macpherson Range) .—247, pseudo-
spretella Sttn. Brit. Tin., p. 14 (Introduced; in Houses). = improbella WIk.,
Char. Lep. Het., p. 86; Turn., Mem. Nat. Mus. Melb., iv, p. 7 (Stanthorpe; moun-
tain areas of N.S.W. and S.A.; Victoria, Tasmania).

BoRKHAUSENIA HEMILEUCA Turn.
The type of this species, which I described in 1896, should be in the South
Australian Museum. I have not recognized it since and think it is improbable
that it really belongs to this genus.

167. BoRKHAUSENIA LEPTOPHYLLA, N. SD.
AerroduAdos, Slender-winged.

2. 183 mm. Head orange-ochreous. Palpi with second joint not reaching base
of antennae, terminal joint three-fifths; ochreous, terminal joint and base of
second joint on outer surface dark fuscous. Antennae fuscous. Thorax fuscous;
apices of tegulae ochreous. Abdomen grey; tuft, sides, and undersurface pale
ochreous. Forewings narrow, elongate, costa nearly straight, apex rounded, termen
very obliquely rounded; orange-ochreous; markings and some irroration dark
fuscous; a moderate basal fascia; an irregular line from one-fourth costa to mid-

BY A. J. TURNER. 91

dorsum, dilated on costa, bent longitudinally and again transversely in disc; a
large costal spot at two-thirds, giving off two lines, one to termination of first
line, the other nearly to dorsum at three-fourths; a similar costal spot at five-
sixths giving off two lines, one joining termination of preceding line, the other to
tornus; cilia orange-ochreous; bases fuscous, Hindwings very narrowly ovate-
lanceolate; fuscous; cilia fuscous.

North Queensland: Eungella (2,500 ft.) in September; one specimen received
from Mr. G. M. Goldfinch, who has the type.

176. BorKHAUSENIA HOMOPELA, Nl. SD.

ouomedos, uniformly dusky.

dg. 17-19 mm. Head and thorax fuscous. Palpi with second joint reaching
base of antennae, terminal joint two-thirds; fuscous. Antennae fuscous; ciliations
in male 1. Abdomen grey-brown. Legs fuscous. Forewings moderately broad,
costa gently arched, apex rounded, termen very obliquely rounded; pale fuscous
without markings; cilia pale fuscous. Hindwings elongate-ovate; whitish-grey;
cilia whitish-grey.

The forewings are much broader and the hindwings much paler than in
B. nyctora Meyr.

Western Australia: Denmark in March; three specimens received from Mr.
W. B. Barnard, who has the type.

182. BoRKHAUSENIA TANYTRICHA, Nl. SDP.
travutpixos, long-haired.

g. 13-14 mm. Head and thorax fuscous-brown. Palpi with second joint not
reaching base of antennae, terminal joint 1; fuscous mixed with ochreous-whitish.
Antennae fuscous-brown with black annulations; ciliations 6. Abdomen brownish-
ochreous. Legs fuscous; tibiae and tarsi with whitish-ochreous rings; posterior
pair mostly whitish-ochreous. Forewings narrow, costa moderately arched, apex
rounded, termen obliquely rounded; pale ochreous-brown finely and closely irrora-
ted with fuscous; four blackish discal spots, first before one-third, second elongate
following first and in a line with first and third, third before two-thirds; pale
spots between first and second, second and third, and beneath third; plical slightly
beyond first discal, a fuscous spot on five-sixths costa; cilia concolorous. Hind-
wings pale fuscous; basal two-fifths whitish-ochreous, tolerably well defined; cilia
pale fuscous, on tornus and dorsum whitish-ochreous.

Queensland: National Park (2,500-3,000 ft.) in November; two specimens.

1838. BoRKHAUSENIA PHANEROSTICTA, 0. SD.

gpavepoorixtos, With conspicuous dots.

do, 9. 18-19 mm. Head whitish-ochreous; erown more or less fuscous.- Palpi
with second joint exceeding base of antennae, terminal joint three-fifths; dark
fuscous, inner surface and some irroration whitish-ochreous. Antennae whitish-
ochreous annulated with blackish; ciliation in male 3. Thorax fuscous-brown.
Abdomen pale brownish. Legs fuscous; tibiae and tarsi ringed with whitish-
ochreous; posterior pair mostly whitish-ochreous. Forewings narrow, oval, costa
moderately arched, apex round-pointed, termen very obliquely rounded; whitish-
ochreous irrorated with fuscous; stigmata blackish, first discal at one-fourth,
second before and third after middle, all in line, plical beyond first discal; a
series of dark fuscous dots near margin from four-fifths costa to a dorsal dot

92 REVISION OF AUSTRALIAN LEPIDOPTERA. OECOPHORIDAE. 1i,

before tornus; cilia whitish-ochreous with a few fuscous points. Hindwings and
cilia pale grey.

Queensland: National Park (4,000 ft.) in November and December; three
specimens.

184. BoRKHAUSENIA PERIGRAPTA, Nl. SD.
Teptypanrtos, Imarked round.

g. 18 mm. Head and thorax pale grey. Palpi with second joint just
reaching base of antennae, terminal joint two-thirds; grey-whitish, external surface
of second joint fuscous towards base. Antennae grey-whitish, towards base
fuscous; ciliations in male 24. Abdomen grey; tuft ochreous-whitish. Legs
fuscous; middle tarsi with whitish rings; posterior pair mostly whitish. Fore-
wings elongate, somewhat dilated posteriorly, costa slightly arched, apex rounded,
termen obliquely rounded; pale grey; markings and some scattered scales blackish;
a moderate sub-basal transverse fascia; first discal at one-third, plical slightly
before it, second discal before two-thirds; a conspicuous series of four costal dots
from middle to near apex, thence a submarginal series of much smaller dots to
tornus; cilia pale grey with a few blackish points. Hindwings elongate-ovate;
whitish-grey; cilia whitish-grey.

Not near any other species.

Western Australia: Denmark in March; one specimen received from Mr.
W. B. Barnard.

185. BoRKHAUSENIA CHALCOTEUCTA, 1. SD.

XadyxoTeukTos, brassy.

fg. 19 mm. Head whitish-ochreous. Palpi with second joint not reaching
base of antennae, terminal joint two-thirds; fuscous, inner surface whitish-
ochreous. Antennae fuscous; ciliations in male 2, Thorax dark fuscous. Abdomen
grey. Legs fuscous (posterior pair missing). Forewings narrow, broadest towards
base, costa rather strongly arched, apex acute, termen extremely oblique; ochreous-
white; markings fuscous; a costal streak throughout, very narrow to one-third,
thence broader; a dorsal streak from one-fourth extended along tornus and termen
to apex; a transverse fascia from one-third costa to mid-dorsum, interrupted
above fold; a second fascia from two-thirds costa to tornus; cilia fuscous, on
tornus grey.

Nearest B. hypochalcha Meyr.

Victoria: Gisborne in January; one specimen. Type in Coll. Lyell.

187. BorKHAUSENIA OXYPEUCES Turn.
I described this as a Hulechria. In the type, 7 of forewings runs to apex, but
in a second example from Victoria to the costal side of apex. In wing-shape and
markings it is allied to B. hypochalcha Meyr.

192. BorKHAUSENIA GYPSOMICTA, N. SDP.
yuwourxtos, Mixed with whitish.

6. 20-21 mm. Head grey; side-tufts whitish. Palpi with second joint just
reaching base of antennae, terminal joint three-fifths; fuscous. Antennae fuscous;
ciliations in male two-thirds. Thorax fuscous; apices of tegulae and a large
central spot whitish. Abdomen dark grey; tuft whitish-ochreous. Legs fuscous;
posterior pair with inner surface of tibiae and tarsal rings whitish-ochreous.
Forewings elongate, narrow, costa gently arched, apex round-pointed, termen
extremely oblique; whitish with dense fuscous irroration, appearing grey; median

BY A. J. TURNER. 93

area more whitish; stigmata dark fuscous, first discal at one-third, plical beneath
it, second discal at two-thirds, an additional dot between the last and tornus; cilia
grey. Hindwings ovate-lanceolate; 3 and 4 coincident; grey; cilia grey.

Queensland: Stanthorpe in December; two specimens received from Mr. W. B.
Barnard, who has the type.

195. BorKHAUSENIA LECHRIOMOCHLA, 0. SD.
Nexprouoxdos, With oblique bars.

6. 24 mm. Head pale grey. Palpi with second joint reaching base of
antennae, terminal joint three-fourths; grey. Antennae fuscous; ciliations in
male 1. Thorax fuscous. Abdomen grey; apices of segments grey-whitish; tuft
whitish-ochreous. Legs fuscous; posterior pair whitish. Forewings narrow, oval,
costa strongly arched, apex round-pointed, termen very obliquely rounded; grey;
markings dark fuscous; an inwardly oblique fascia from one-third costa to one-
third dorsum; a second fascia from two-thirds costa to termen above tornus,
containing a pale central line; a median discal spot between fasciae; an apical
suffusion; cilia fuscous. Hindwings and cilia brown-whitish.

New South Wales: Barrington Tops, in February; one specimen received
from Mr. G. M. Goldfinch, who has the type.

197. BoRKHAUSENIA NIGRIPUNCTA, 0. SDP.

nigripunctus, black-spotted.

®. 18 mm. Head, thorax, and abdomen grey. Palpi with second joint
reaching base of antennae, terminal joint three-fourths; whitish, terminal joint
grey. Antennae grey. Legs grey. Forewings elongate, somewhat dilated
posteriorly, costa slightly arched, apex pointed, termen very obliquely rounded;
grey-whitish with slight blackish irroration; spots blackish, one on costa near
base, another median, sub-basal, first discal at one-third, longitudinally elongate,
plical obsolete, second discal before two-thirds, a spot above dorsum at three-
fourths; some blackish irroration on and before termen; cilia grey-whitish
irrorated with blackish. Hindwings elongate ovate; pale grey; cilia pale grey.

Tasmania: Cradle Mt. (3,000 ft.) in January; one specimen.

200. BorRKHAUSENIA ALBIPECTINATA, N. SD.
albipectinatus, with white pecten.

6. 17-18 mm. Head grey; margins of crown white. Palpi with second joint
just reaching base of antennae, terminal joint three-fourths; fuscous. Antennae
grey; pecten white; ciliations in male 1. Thorax grey or fuscous irrorated with
whitish. Abdomen grey or fuscous. Legs grey. Forewings narrow, elongate, costa
slightly arched, apex acute, termen extremely oblique; fuscous-grey with fine
whitish irroration; stigmata dark grey, first discal at one-third, plical beneath
it, second discal at two-thirds, cilia grey. Hindwings broadly ovate-lanceolate;
grey: cilia grey.

An obscure species, but the white pecten contrasting with the grey face is
distinctive.

Queensland: Brisbane in October; Toowoomba in November. Two specimens.

202. BoRKHAUSENIA NUBIFERA Meyr.

I have examined the type of B. cyclozona Low., of which the head, abdomen,
and one forewing are now missing. The markings of the forewings are accurately
described, except that the transverse marking is said to be at two-thirds instead
of one-third, a clerical error. The neuration of the hindwings is that of this
genus and not Paratheta.

94 REVISION OF AUSTRALIAN LEPIDOPTERA. OECOPHORIDAE. ii,

203. BoRKHAUSENIA SERRULIFERA, Nl. SD.
serruliferus, with a small saw.

g. 18-19 mm. Head and thorax dark grey. Palpi with second joint just
reaching base of antennae, terminal joint three-fifths, grey. Antennae grey;
ciliations 2. Abdomen grey, posteriorly brownish in dorsum; tuft pale grey.
Legs grey; posterior pair grey-whitish. Forewings elongate-oval, costa moderately
arched, apex pointed, termen very obliquely rounded; grey; a curved, oblique,
finely serrulate, blackish interrupted line from one-third costa to one-third dorsum;
a blackish dot in dise above middle closely following this; some dark fuscous
irroration on tornus and in terminal area; cilia grey. Hindwings ovate; pale
grey; cilia pale grey.

In one example 3 and 4 of hindwings are stalked, in the other connate.

Queensland, in July and August; two specimens.

205. BoRKHAUSENIA IULOPHYLLA, 0. SD.

ivAodvAXos, With softly hairy wings.

do. 20 mm. Head ochreous-whitish. Palpi with second joint not reaching
base of antennae, terminal joint four-fifths, rather stout; dark grey. Antennae
dark grey; ciliations 1. Thorax grey. Abdomen ochreous-grey. Legs grey;
posterior tibiae pale ochreous. Forewings elongate-oval, costa moderately arched,
apex round-pointed, termen rounded, very oblique; pale grey with some fuscous
irroration in terminal area; stigmata fuscous, first at two-fifths, second at two-
thirds, plical before first discal, connected by some fuscous scales with dorsum;
cilia whitish-grey. Hindwings and cilia pale ochreous; a dense crest of long
spreading ochreous hairs from base above.

This is certainly a near ally of Disselia aleurota Meyr., but that genus cannot,
I think, be maintained. The hairy crest which arises from the dorsum of the
second anal vein of the male of both species is merely an exaggeration of a
smaller pencil of hairs rising from near the base of that vein in the female of
aleurota, and in both sexes of pseudospretella and other species of Borkhausenia.

Western Australia: Albany in February; one specimen in Coll. Barnard.

208. BoRKHAUSENIA GYPSODES, N. Sp.
yuywdns, chalk-white.

6. 18 mm. Head grey. Palpi with second joint not reaching base ot
antennae, terminal joint three-fifths; fuscous. Antennae fuscous; ciliations 2.
Thorax fuscous, margins suffusedly pale grey. Abdomen grey, dorsum except
base fuscous-brown; tuft whitish-ochreous. Legs fuscous; posterior pair ochreous-
whitish. Forewings suboval, costa gently arched, apex rounded, termen very
obliquely rounded; whitish, margins suffused with grey; stigmata fuscous, first
discal at one-third, plical well before it; second discal at two-thirds; a sub-
terminal series of small fuscous dots from beneath five-sixths costa to tornus:
pale grey. Hindwings ovate; pale grey; cilia whitish.

Western Australia: Kalamunda, near Perth, in January; one specimen
received from Mr. W. B. Barnard.

210. BoRKHAUSENIA MISELLA, 0. Sp.
misellus, miserable.
fg. 18 mm. Head whitish-ochreous. Palpi with second joint not reaching base
of antennae, terminal joint three-fourths; fuscous, internal surface and terminal
joint whitish-ochreous. Antennae grey; ciliations in male two-thirds. Thorax

BY A. J. TURNER. 95

fuscous. Abdomen grey. Legs fuscous; posterior pair whitish-ochreous. Fore-
wings elongate-oval, costa moderately arched, apex pointed, termen very oblique;
grey; markings fuscous; a subcostal dot at one-sixth; a thick outwardly oblique
line from beneath one-fourth costa to fold, there curved outwards to become
longitudinal for a short distance, both ends rounded, second discal at three-
fifths, with an additional dot below and beneath it; apical area suffused with
fuscous; cilia grey. Hindwings elongate-ovate; pale grey; cilia pale grey.

Queensland: National Park (3,000 ft.) in November; two specimens received
from Mr. W. B. Barnard, who has the type.

212. BorKHAUSENIA VERNILIS, D. Sp.

vernilis, mean. ;

fg. 18-19 mm. Head, thorax, and abdomen fuscous. Palpi with second
joint reaching base of antennae, terminal joint three-fifths; fuscous. Antennae
fuscous; ciliations in male 1. Legs fuscous. Forewings elongate-oval, costa
moderately arched, apex pointed, termen very obliquely rounded; fuscous with a
few dark fuscous scales mostly in terminal area; stigmata small or nearly
obsolete, dark fuscous, first discal at one-fourth, plical beneath it, second discal
slightly beyond middle, sometimes an additional dot between and above discals;
cilia fuscous. Hindwings elongate-ovate; fuscous; cilia fuscous.

Western Australia: Denmark in March and April; four specimens received
from Mr. W. B. Barnard, who has the type.

213. BoORKHAUSENIA CNECOCRANA, DL. SD.

xynkoxpavos, With yellowish head.

cd. 18-20 mm. Head whitish-ochreous. Palpi with second joint not reaching
base of antennae, terminal joint two-thirds; dark fuscous. Antennae fuscous;
ciliations in male two-thirds. Thorax fuscous. Abdomen grey; tuft ochreous-
whitish. Legs fuscous; posterior pair grey. Forewings elongate, costa slightly
arched, apex pointed, termen very oblique; grey-whitish; irrorated with fuscous
except in middle and towards base; a thick fuscous streak on costa to two-
thirds; stigmata dark fuscous, first discal at one-third, confluent with costal
streak, plical slightly beyond it, second discal at two-thirds, additional dots
above and below middle, and beneath second discal; cilia grey. Hindwings
elongate-ovate; grey; cilia grey.

New South Wales: Sydney, in October; two specimens received from Mr.
G. M. Goldfinch, who has the type.

214. BoRKHAUSENIA FLAVIPUNCTA, 0. SD.
fiavipunctus, yellow-spotted.

6. 12 mm. Head white. Palpi with second joint much exceeding base of
antennae, terminal joint three-fifths; second joint whitish, basal half of outer
surface and a subapical ring fuscous, third joint fuscous. Antennae fuscous;
ciliations in male 1. Thorax fuscous. (Abdomen missing.) Legs ochreous-
whitish; anterior pair fuscous. Forewings narrow, costa gently arched, apex
rounded, termen very obliquely rounded; white with some fuscous irroration; a
suffused yellow spot on base of dorsum; stigmata fuscous, first discal at one-
third, plical before it, second discal before two-thirds; between these is a broad
irrorated band, and apical area is also irrorated; cilia pale fuscous, a spot on
tornus yellow. Hindwings elongate-ovate; narrow; grey; cilia grey.

North Queensland: Kuranda in August; one specimen.

96 REVISION OF AUSTRALIAN LEPIDOPTERA. OECOPHORIDAE. il,

216. BoRKHAUSENIA LISSOPTERA, ND. SDP.

\uscomrepos, SMooth-winged.

gd, 9. 12-16 mm. Head and thorax ochreous-whitish. Palpi with second
joint reaching base of antennae, terminal joint three-fifths; second joint with
median and subapical, terminal joint with basal and apical pale fuscous rings.
Antennae ochreous-whitish annulated with pale fuscous; ciliations in male two-
thirds. Abdomen grey. Legs fuscous annulated with ochreous-whitish; posterior
pair ochreous-whitish. Forewings sub-oval, costa moderately arched, apex round-
pointed, termen very obliquely rounded; ochreous-whitish with slight fuscous
irroration, mostly in terminal half; stigmata fuscous, first discal at one-third,
plical before it, second discal before two-thirds; a series of fuscous dots close to
termen and apical half of costa; cilia ochreous-whitish, on tornus more or less
fuscous. Hindwings elongate-ovate; pale grey; cilia ochreous-grey-whitish.

North Queensland: Kuranda in September; Queensland: Stradbroke J. in
September among coastal jungle. Five specimens.

217. BoRKHAUSENIA LEPTOCNECA, Nl. SD.

Aertoxvynkos, Slightly yellowish.

gd. 16-18 mm. Head yellow-whitish. Palpi with second joint much exceeding
base of antennae, terminal joint one-half; whitish, basal two-thirds of external
surface of second joint fuscous, second joint with subapical, terminal joint with
basal and broad apical fuscous rings. Antennae fuscous; ciliations in male 1.
Thorax fuscous; anterior edge and a posterior spot whitish. Abdomen grey;
tuft ochreous-whitish. Legs ochreous-whitish; anterior pair fuscous. Forewings
dilated posteriorly, costa moderately arched, apex pointed, termen nearly straight,
oblique; whitish, towards dorsum yellowish-tinged, rather densely irrorated with
pale fuscous except on dorsum; markings pale fuscous; a broad costal streak
from base to one-fourth; an oblique line from one-fourth costa to dorsum near
middle; sometimes interrupted above dorsum; first discal at one-third, connected
with this line; second at two-thirds connected by a line with tornus; an inter-
rupted line or series of dots close to termen and apical third of costa; cilia
grey, on tornus pale yellowish. Hindwings elongate-ovate; pale grey; cilia
pale grey.

North Queensland: HEungella (2,500 ft.) in September; three specimens
received from Mr. G. M. Goldfinch, who has the type.

224. BorRKHAUSENIA BASILEUCA, n. sp.
Baovrevxos, whitish at the base.
3d. 18 mm. Head and thorax dark fuscous; apices of tegulae whitish. Palpi

with second joint reaching base of antennae, terminal joint 1; dark fuscous.’

Antennae dark fuscous; ciliations in male 13. Abdomen grey; tuft ochreous-
whitish. Forewings narrow, costa gently arched, apex pointed, termen very
oblique; fuscous; stigmata obscure, dark fuscous, first discal at one-third, plical
beneath it, second discal at two-thirds; a small but distinct median whitish spot
at base; cilia fuscous, apices grey. Hindwings and cilia grey. :

Very obscure, but should be recognizable by the white spots on apices of
tegulae and bases of forewings.

Western Australia: Mt. Dale in January; one specimen received from My.
W. B. Barnard.

BY A. J. TURNER. 97

225. BoORKHAUSENIA LECHRIOGRAMMA, Nl. SDP.
Aexptoypaumos, Obliquely marked.

9. 18-20 mm. Head and thorax whitish irrorated with grey. Palpi with
second joint exceeding base of antennae, terminal joint three-fifths; whitish
partly suffused with grey. Antennae grey. Abdomen whitish-grey; bases of
segments on dorsum ochreous-brown. Legs grey; posterior pair ochreous-whitish.
Forewings moderately broad, slightly dilated posteriorly, costa moderately arched,
apex pointed, termen slightly rounded, oblique; whitish uniformly irrorated with
grey and patchily with fuscous; markings fuscous; first discal at one-third, plical
beyond it, elongate, second discal at two-thirds, included in a line from three-
fifths costa to termen above tornus; a line from four-fifths costa, sharply angled
before apex, and continued close to termen to tornus; cilia grey-whitish. Hind-
wings elongate-ovate; pale grey; cilia grey-whitish.

Western Australia: Busselton in October; two specimens received from Mr.
G. M. Goldfinch.

238. BoRKHAUSENIA PSARITIS, N. SD.
Wapiris, ashen-grey.

6, . 16-19 mm. Head white. Palpi white; second joint exceeding base
of antennae, terminal joint three-fifths; second joint with basal half and
a subapical ring fuscous; terminal joint sometimes partly fuscous. Antennae
dark fuscous; ciliations in male one-half. Thorax dark fuscous. Abdomen fuscous.
Legs whitish; anterior pair mostly fuscous; middle tibiae and tarsi annulated
with fuscous. Forewings rather narrow, suboblong, costa rather strongly arched,
apex round-pointed, termen obliquely rounded; white with fuscous markings; a
narrow basal fascia prolonged as a broad streak along costa to one-fourth; an
inwardly oblique fascia from one-third costa to one-fourth dorsum, anteriorly
well defined, and with a median darker dot, posteriorly broadly suffused on costa
and dorsum; a second fascia confluent with preceding on costa to tornus, joined
by a broad streak from apex; posterior area of disc is mainly fuscous, but there
are three white patches of variable size left by these fasciae, one dorsal, one
costal, one terminal; an interrupted terminal line; cilia fuscous-whitish. Hind-
wings and cilia pale grey.

Queensland: National Park (3,000 to 3,500 ft.) in December, January and
March; 23 specimens.

241. BoRKHAUSENIA PELOPHANES, N. SD.

medoparyns, dusky.

dg. 18 mm. Head and thorax fuscous-brown. Palpi with secend joint just
reaching base of antennae, terminal joint three-fourths; brownish, outer surface
of second joint irrorated with blackish, base of terminal joint blackish. Antennae
fuscous-brown; in male serrate, ciliations in male one-half. Abdomen grey; tuft
whitish-ochreous. Legs fuscous with whitish-ochreous rings; posterior pair
mostly whitish-ochreous. Forewings narrow, posteriorly dilated, costa gently
arched, apex rounded, termen obliquely rounded; fuscous-brown; stigmata blackish,
first discal at one-third, plical before it, second discal slightly beyond middle;
cilia brown. Hindwings and cilia grey.

Tasmania: Russell Falls in January; one specimen.

98 REVISION OF AUSTRALIAN LEPIDOPTERA. OECOPHORIDAE. iil.

244. BoORKHAUSENIA CENTROSTICHA, Nl. SD.

KevTpoortxos, With central streak.

dg. 24mm. Head and thorax fuscous. Palpi with second joint reaching base
of antennae, terminal joint one-half; fuscous with some whitish scales, apex of
second joint whitish. (Abdomen missing.) Legs fuscous; tarsi with whitish
rings. Forewings elongate, somewhat dilated, costa gently arched, apex round-
pointed, termen straight, oblique; fuscous; a blackish median streak from one-
third to nearly two-thirds; its anterior end bent slightly upwards, followed
posteriorly by a blackish dot, both partly edged with whitish; cilia fuscous,
apices paler. Hindwings rather broadly ovate; pale grey; cilia pale grey.

Very sombre, but very distinct by the curious central streak of forewings.

Victoria: Gisborne in August; one specimen from pupa found under bark
(R. W. Hill). Type in Coll. Lyell.

245. BoRKHAUSENIA NEPHOTYPA, D. Sp.
vepoturos, Cloud-marked.

9. 24 mm. Head and thorax grey. Palpi with second joint reaching base of
antennae, terminal joint four-sixths, grey-whitish with a few fuscous scales,
terminal joint fuscous. Antennae grey. Abdomen whitish-grey. Legs grey;
posterior pair whitish. Forewings elongate-oval, costa rather strongly arched,
apex round-pointed, termen nearly straight, strongly oblique; grey; a strong
undefined dorsal suffusion cut by an inwardly oblique grey-whitish streak above
one-fifth dorsum; first discal at one-third, plical before it, confluent with dorsal
suffusion, second discal before two-thirds; cilia grey. Hindwings and cilia
pale grey.

New South Wales: Waterfall near Bulli in September; one specimen received
from Mr. G. M. Goldfinch, who has the type.

246. BorKHAUSENIA TRICHOCEROS, N. SD.
Tpikoxepws, With hairy horns.

6. 18-19 mm. Head and thorax ochreous-fuscous. Palpi with second joint
reaching base of antennae, terminal joint three-fifths; whitish-ochreous irrorated
with fuscous. Antennae ochreous-whitish annulated with pale fuscous; ciliations
in male 5. Abdomen whitish-ochreous. Legs fuscous; posterior pair whitish-
ochreous. Forewings suboval, costa moderately arched, apex round-pointed,
termen very obliquely rounded; whitish-ochreous with slight patchy fuscous
irroration; first discal at one-fourth, plical beyond it, second discal before two-
thirds, an additional spot midway between first and second, all dark fuscous; an
indistinct series of fine fuscous dots close to termen and apex; cilia whitish-
ochreous with a few fuscous points. Hindwings elongate-oval; grey-whitish;
cilia ochreous-whitish.

There is a pencil of long hairs arising from the dorsum of the second anal
vein of the hindwings just as in B. pseudospretella and some other species.

Queensland: National Park (4,000 ft.) in November; two specimens.

ON THE PRODUCTION OF FERTILE HYBRIDS FROM CROSSES BETWEEN
VULGARE AND KHAPLI EMMER WHEATS.
By W. L. WaTrernHousE, The University of Sydney.
(Plates ili—iv.)
[Read 26th April, 19383.
Introduction.

Crosses between common wheats (Triticum vulgare L.) having 21 pairs of
chromosomes and other species having 14 pairs can sometimes be made with
comparative ease, but extreme difficulty is encountered in other cases. Some
members of the emmer group (7. dicoccum Schrnk.) fall within this latter
category. An outstanding example is the Indian emmer known as ‘“Khapli’.
Botanically it is a member of the group 7. dicoccum Ajar Pere. From a
commercial wheat-growing point of view it is unimportant, but it is a remarkable
wheat for its extreme resistance to disease. There are few varieties showing
this quality to the same degree.

On account of its disease resistance, many efforts have been made to cross
“Khapli” with vulgare wheats. Puttick (1921) found that crosses with “Marquis”
(T. vulgare) gave only sterile plants. Hayes and Stakman (1922) reported
disappointing results from attempts to make the same crosses. Hynes (1926)
described work giving fertile F, plants from crosses between “Federation” and
“Khapli’”. Thompson and Hollingshead (1927) secured grain from crosses with
vulgare wheat, but no fertile plants resulted. Waterhouse (1930) reported
similar failure in an extensive series of crosses between vulgare wheats and
“Khapli”. Hollingshead (1932) failed when using “Federation”, but met with
a measure of success from using as the vulgare parent a wheat which had been
produced by McFadden (1930) from a cross between a vulgare and an emmer
wheat.

There is a further record which does not appear to have been published.
in communications to the writer under date 22nd September, 1929, and 23rd
September, 1930, the Economic Botanist of the Central Provinces of India has
stated that certain vulgare wheats in cultivation there have been derived from
crosses between “Murya’, a common wheat, and ‘‘Khapli’ emmer. He kindly
forwarded grain of these wheats. They have been grown and it is found: that
this “Khapli’” is totally different from the ‘Khapli’ obtained from Dr. E. C.
Stakman of Minnesota under the C.Il. Number 4013, the strain used by the
other workers. It falls within a different subspecies, having pubescent chaff and
black awns.

Obviously this Indian crossing work involves a totally different parent. It
may be remarked that crosses between “Federation” and this emmer have given
sterile F, plants similar to those reported for the true “Khapli” crosses. The
same result has been obtained when “Murya’ has been crossed with the true
“Khapli”.

100 CROSSES BETWEEN VULGARE AND KHAPLI EMMER WHEATS,

Further “Khapli” Crosses.

Since the summarizing of the writer’s previous work (Waterhouse, 1930)
up to the 1927 season, further ‘‘Khapli”’ crosses have been made each year. A
recent striking success makes it wise to report the results at this stage, although
the investigations are still in progress.

Having consideration to the possible effect that seasonal variations may
have upon the results of crossing, certain vulgare parents—and notably
“Federation’—have been used year after year. Whilst there have been some
differences in the number of grains set, in their plumpness and in the amount
of growth made, the characters of the F, plants have not appreciably varied
from season to season. In all cases they have been completely sterile ‘grass
clumps” when “Federation” has been used.

The results to date in respect to the grain setting from the crosses made
are summarized in Table 1.

TABLE 1.

Summary of results obtained from crosses between vulgare and ‘“Khapli”’ wheats.

Number of Number of Percentage
Year of Crosses. Grains Set. Flowers Pollinated. Grain-Setting.
*1921 to 1927 .. se Bis Sha ne 286 1,860 15
Ie yssy) Gis we a ae A a0 29 118 25
1929) ee as us ou ae 197 654 30
19380... ia ae oH he ef 64 136 48
OST seus a Ge a Bm 633 1,278 50
1932" sae Be ee ae ee ys 618 1,134 54
Totals .. Ate ue ao 1,827 5,180

* Vide Proc. LINN. Soc. N.S.W., 55, Part 5, p. 605.

The average grain-setting for the period amounts to 35%. In the earlier
years “Federation” and “Hard Federation’ were extensively used, but more
recently only 1 or 2 heads of these varieties have been pollinated each season.
The increase in the grain-setting in later years is mainly due to the use of other
vulgare wheats which cross somewhat more readily with “Khapli”.

The varieties of vulgare wheat which have been used to date in the crosses
with ‘“‘Khapli” are as follows:

Akakomoughi Etawah (Indian F x Telford’s)
Anchor Exquisite Kenya Crossbred C6041
Aussie Federation e 34 C6042

Baroota Wonder

- Bearded Gluyas

(Federation x Khapli)
Felix

Kenya Governor
King’s Early

Bena Firbank Linden

Bobin Florence Little Club

Bobs Ford Mac’s White
Bomen Free Gallipoli Maharajah

Bunge Gallipoli Marquillo

(Bunge X Emmer 19) Gatra Murya

Bunyip Geeralying (Murya x Khapli)

BY W. L. WATERHOUSE.

101

Canberra Gem Nabawa
Cedar Gluyas No. 76
Clarendon Gresley Penny
Clubhead Guinea (Purple Straw x Medeah)
Comeback Gullen Quantity
Crossbred W519 Hard Federation Queen Fan
. W522 Hope Riverina
Currawa Hornbill (Stanley x Yandilla King)
Dindiloa (Huguenot x Federation) Steinwedel
Dundee (Huguenot x Fed’tion « Ied’tion) Sunset
Duri Hurst’s 11 Thew
Early Bird Improved Steinwedel Waratah

Early Defiance Indian F Yandilla King
(Emmer X Marquis) Indian 12 Zaft

Certain of these crosses were actually made by Messrs. J. H. Kaye and
J. Bolin, whose help has been loyally given out of their own time and is grate-
fully acknowledged. The first of the Steinwedel and Improved Steinwedel crosses
were actually made by the former.

It was early considered that vulgare wheats owing their origin to crosses
involving wheats with 14 pairs of chromosomes might cross. with ‘‘Khapli’? more
readily than other vulgares. All available wheats of this nature have been tried,
as the above list shows.

The results have been extremely variable. The crossed grain in some cases
has been tiny and pinched, in others large and plump. The grain has failed to
germinate in some instances. In others the F, plants have been tiny “grass
clumps” which have soon died. All the intermediate stages have been found
between these and fairly well grown plants up to 8 feet high and showing a
low degree of fertility. With the three exceptions to be described presently, all
have shown notable stunting and yellowing, followed by early death, even under
conditions of careful cultivation and watering. This extreme chlorosis has been
a striking feature.

Seedling tests of these F, plants have been made with three physiologic
forms of Puccinia graminis tritici. With Forms 43 and 46 there has been an
approach to complete dominance of resistance. With Form 34 the approach has
been to dominance of susceptibility. Flag smut tests have shown dominance of
resistance.

The vulgare parents of the crosses in which the F, plants have produced a
grain or grains are as follows: Anchor, Bunyip, Canberra, Crossbred W 519,
Exquisite, Florence, Free Gallipoli, Garra, Geeralying, Gresley, Gullen, Hornbill,
Improved Steinwedel, Linden, Marquillo, Riverina, Steinwedel, Zaff.

These include one or two cases in which pollen of the vulgare parent was
used to pollinate the F, stigmas, in addition to those in which open-pollination
took place.

An examination of the available pedigrees of these varieties has been made.
It shows that seven out of the eighteen which gave fertile progeny are derivatives
cf crosses involving T. durum. On the other hand it is found that ten of the
sixty parents which produced sterile F, plants were durum or dicoccum derivatives,
including a “fixed” vulgare wheat from the (Federation x Khapli) cross reported
by Hynes, and “Hope”, a dicoccum derivative.

From certain of the crosses—notably when “Bobin”, “Gullen” and “Geeralying’”’
are the vulgare parents—material is now in the F, generation. Extremely wide
segregation is being shown. Analysis is not complete, but seemingly there is a
paucity of vulgare types.

202 CROSSES BETWEEN VULGARE AND KHAPLI EMMER WHEATS.

The grain-setting by the F, plants under conditions of open-pollination is low.
Thus in 1932, which was a good season, 153 grains were produced from 224 F,
plants of 11 different vulgare and ‘‘Khapli” crosses.

It will be seen that the fertility of these crosses has been low and the
advance made in the problem not very great.

Higher Fertility Crosses.

In addition to considering the likelihood of durum or dicoccum parentage in
vulgare wheats contributing to ease of crossing, those varieties like “Bobin”’
and “Gullen” which gave a small measure of fertility were studied in regard
to their parentage. Where possible the ancestors of these wheats were used
in further crosses with “Khapli”’, together with other varieties in whose pedigrees
these same ancestors appeared. Of course there were others in which there was
merely haphazard selection of the vulgare parent. But the success gained is trace-
able to the choice of wheats entering into the pedigree of those which had earlier
given some slight fertility.

In 1931 “Garra’, “Improved Steinwedel”, and “Steinwedel’ were pollinated
with “Khapli” pollen. Unusually plump grain was set to the extent of about
65%, which was approximately the average grain-setting in the intra-species
crosses made that season. Full germination was obtained and the F, seedlings
were so healthy and vigorous as almost to make one doubt their hybridity. Their
rust reactions, however, showed them to be certainly crosses. The flag smut
tests were equally conclusive. After transplanting to the open field, these cross-
breds showed astonishing vigour in the midst of many other crossbreds which
were weak and yellow. At maturity the plants were normal in development
(Plate iii). The leaves were dark green, tillering abundant and heads of
normal size. <A general intermediacy of characters in relation to the parents
was exhibited.

Heads of each of the three crosses were bagged to ensure self-pollination.
A little less than 1% of grain set in these heads. Back-crosses with the vulgare
parents gave a Slightly higher fertility and this was still higher when the F,
plants were pollinated with “Bobin” and ‘Florence’,

Most of the heads were open-pollinated. The results from the harvest are
shown in Table 2, and typical heads are illustrated in Plate iv.

TABLE 2.
Grain setting in crosses between three vulgare wheats and “Khapli’” emmer.

|

Number of Number of Number of Percentage
Parents of Cross. F, Plants Grains Flowers Grain-
Counted. Set. Present. | Setting.
Garra X Khapli_.. at is ae 15 ASS 8,769 13°2
Improved Steinwedel x Khapli .. bes Tat 1,105 4,522 | 22-2
Steinwedel x Khapli 5H ae ES 13 1,307 6,804 19-1
Totals fe = co | 39 3,565 if 20,095
|

BY W. L. WATERHOUSE. 103

There is an average grain-setting of 17-2%. This grain varies from shrivelled
to normal plump grain, as shown in Plate iv.

When compared with the weakly plants and insignificant fertility obtained by
the use of other vulgare parents, this is a remarkable result.

In the 1932 season the same crosses were repeated, together with a number
of others in which the vulgare parent has in its pedigree one or other of the three
wheats used so successfully. Unusually good grain-setting has been obtained
from some of these.

It seems probable that this capacity of the three vulgare wheats to cross
with ‘“Khapli” will have a wider application. There are other refractory wheats
in addition to “Khapli”, having 14 pairs of chromosomes and crossing only with
difficulty (if at all) with “Federation” and other vulgare wheats. As examples,
“Gaza’’, “Beladi’, and “Iumillo”’ might be cited. In 1932 these were crossed
with “Garra’’, “Improved Steinwedel’, and ‘“Steinwedel’, and an unusually high
setting of plump grain has been obtained. In crosses with rye, however, the
same success has not been maintained.

The three vulgare wheats are related. “Garra” comes from a cross involving
“Steinwedel’’, as does also “Improved Steinwedel”. The other wheats which
enter into their pedigrees give sterile, or almost sterile F, plants when crossed with
“Khapli”’. “Steinwedel” is a selection from “Champlain’s Hybrid’. Efforts have
been made to secure a sample of this wheat, but have failed. Cytological studies
are therefore planned using “Steinwedel” as a basis.

Acknowledgments.

In addition to acknowledgments included in the text, thanks are due to
Wire, dig ak Pridham, of Cowra, who originally supplied grain of the wheats
reported on. Financial assistance from the Trustees of the Science and Industry
Endowment Fund is gratefully acknowledged.

Summary.

More than 5,000 crosses between vulgare wheats and ‘“‘Khapli”’ emmer have
been made. An average of about 35% grain-setting has resulted. Almost always
sterility has been shown by the progeny, but in a few cases a low measure of
fertility has been found. Some of these derivatives are now in the F, generation.

A big advance comes from crosses made in 1931. The vulgare wheats known
as “Garra’’, “Improved Steinwedel” and “Steinwedel”’ are found to cross readily
with “Khapli” and have given F, plants of normal development. Under conditions
of open-pollination, these hybrid plants have set more than 17% of grain. Crosses
between these three vulgare wheats and other refractory members of other species
of Triticum indicate an unusually high fertility in these cases also. Of the three
wheats, “Steinwedel” is the one to which this capacity for crossing is traceable.

Literature Cited.

HAyYeEs, H. K., and STakMaNn, EH. C., 1922.—Wheat stem rust from the standpoint of
plant breeding. Proc. Second Ann. Mtg. Canadian Soc. Agron., 1921, pp. 997-1012.

HOLLINGSHEAD, L., 1932.—Partly fertile hybrids of common wheat with Khapli emmer.
Journ. Heredity, 23, pp. 247-253.

HYNEs, H. J., 1926.—Studies on the reaction to stem rust in a cross between Federation
Wheat and Khapli emmer, with notes on the fertility of the hybrid types. Phyto-
pathology, 16, pp. 809-827.

McFabDpDEN, E. S., 1930.—A successful transfer of emmer characters to vulgare wheat.
Journ. Amer. Soc. Agron., 22, pp. 1020-1034.

104 CROSSES BETWEEN: VULGARE AND KHAPLI EMMER WHEATS.

PuTrTick, G. F., 1921.—The reaction of the F, generation of a cross between a common
and a durum wheat to two biologie forms of Puccinia graminis. Phytopathology, 11,
pp. 205-2138.

THOMPSON, W. P., and HouuINGsHEAD, L., 1927.—Preponderance of dicoccum-like charac-
ters and chromosome numbers in hybrids between Triticwm dicocewm and T. vulgare.
Journ. Genetics, 17, pp. 283-307.

WATERHOUSE, W. L., 1930.—Australian Rust Studies, III. Initial results of breeding for
rust resistance. Proc. LINN. Soc. N.S.W., 55, pp. 596-636.

EXPLANATION OF PLATES III-IV.
Plate iii.

A.—Group of plants of (Steinwedel x Khapli) at flowering time. Some of the
strongly chlorotic and stunted plants of other sterile crosses may be seen in the fore-
ground. Scale in feet at right of picture.

B-F.—Single plants of wheats grown under similar conditions and harvested at
maturity. B.—‘‘Khapli’’. C.—“Steinwedel”’. D.—Fertile F, plant of (Steinwedel
Kkhapli). E.—Fertile F, plant of (Improved Steinwedel x Khapli). F.—Fertile F, plant
of (Garra x Khapli). Scale next to fig. C is one metre long.

G.—Grain produced by one of the F, plants of (Garra x Khapli). Natural size.

Plate iv.
A-G.—Pairs of heads of typical wheat plants grown under similar conditions and
shown half natural size. A.—‘‘Khapli’. B.—‘“Steinwedel’. C.—‘‘Improved Steinwedel”’.

D.—“Garra”. E.—F, plant of (Steinwedel x Khapli). F.—F, plant of (Improved
Steinwedel x Khapli). G.—F, plant of (Garra x Khapli).

H-N.—Two pairs of glumes from contiguous spikelets of wheat heads. Natural
size. H.—‘‘Khapli’’. I.—‘‘Steinwedel”’, J.—‘Improved Steinwedel’’. K.—“‘Garra’”’.
L.—F, plant of (Steinwedel x Khapli). M.—F, plant of (Improved Steinwedel x Khapli).
N.—F, plant of (Garra x Khapli).

O-Q.—Grain produced by pollinating three vulgare wheats with pollen of ‘“Khapli”.
Natural size. O.—(Steinwedel x Khapli). P.—(Improved Steinwedel x Khapli). Q.—
(Garra x Khapli).

PLATE IIT.

1933.

N.S.W.,

Proc, Linn. Soc.

ae

PLATE Iv.

INESaVe; lio:

Proc, Linn. Soc.

esi

SOME CLIMATOLOGICAL ASPECTS OF ARIDITY IN THEIR
APPLICATION TO AUSTRALIA.

By Joun AnprEws, B.A., Demonstrator in Geography, University of Sydney,
and W. H. Mazr, Caird Scholar in Geography.
(Seven Text-figures. )
[Read 26th April, 1933.]
Introduction.

This paper deals with the question of the climatological factors involved in
the concept of aridity, and endeavours to apply these to Australian conditions.
In particular, the Index of Aridity as phrased by de Martonne (1927) is examined
in detail for annual conditions in Australia, and the idea extended to a survey
of the monthly conditions. So far as the authors are aware, none of these, except
generalized versions of Figures 1 and 2, has appeared before. In addition to
considering the characteristic appearance of drainage systems, the map and data
have been critically examined with reference to agricultural facts. 'The authors
believe that an attempt should be made to define the nature of aridity, however
complex this may be.

S00 MILES

ANNUAL. 50 ee GF,

40 |

35

is 120 ’

Fig. 1.—The index of aridity for mean annual conditions in Australia. The more arid
the region the lower is the index figure.

SOME CLIMATOLOGICAL ASPECTS OF ARIDITY,

106

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BY J. ANDREWS AND W. H. MAZE. 107

Definitions of Aridity.

Although many writers have discussed arid lands, most have been content to
apply the terms “desert”, “arid”, “semi-arid”, etc., without definitions. The popular
acceptance of the underlying idea has been so complete as to make more explicit
statements unnecessary in non-scientific works. Within the last few decades,
however, when attention became focussed on scientific climatology, attempts have
been made to define the nature of aridity.

These attempts fall into two classes: firstly, those with an empirical basis
and, secondly, numerical definitions which have been made for the purpose of
illustrating climate distributions of continental dimensions. Of the former
some of the more important are as follows:

“A desert is a region of sparse and specialized plant and animal life”
(Isaiah Bowman) ;

“A desert is a region with a rainfall of 10 ins. or less per annum, where
agriculture is impracticable, and where occupation is found possible only for
a sparse population of pastoralists” (J. W. Gregory) ;

“A desert is a region of small rainfall (sometimes, however, amounting
to 16 ins. per annum in a hot region) with a sparse and specialized plant and
animal life. It is not found capable of utilization by stationary pastoralists
even after the borders have been occupied by this class for 50 years” (Griffith
Taylor).

These definitions of course leave room for very considerable variation in the
drawing of boundary lines.

Among the numerical definitions of the second class may be mentioned that
of Mark Jefferson (1916), where he states that in the United States less than
5 inches per annum gives desertic conditions; from 5 to 10 inches gives arid
conditions, and from 10 to 20 inches gives semi-arid conditions. Koppen (1923)
gives 25 cm. (10 in.) as indicative of the desert boundary, and 25 to 50 cm.
(10 to 20 in.) as indicating “steppe” conditions. De Martonne also recognizes
25 em. as significant in the delimitation of desert climates.

These estimates are first approximations, and must be studied in their local
applications. Russell (1931) has recently modified Koppen’s formulae in their
application to dry regions of the United States as a result of field examination
of his climatic boundaries.

Criteria available.

Several points of reference may be made in defining “aridity”, e.g., climato-
logical, physiographic, vegetational, or developmental and occupational. It is
assumed in this paper that a complete definition of aridity must contain reference
to each of these. From our present point of view, however, there are some
considerable difficulties in the way of using any of them for purposes of definition.
The stations taking complete records even of precipitation and temperature alone
are very few in relevant regions in Australia. Again, there is no adequate basis
so far proposed that will enable one to compare in a quantitative manner land-
scapes in various regions. Few studies, too, with notable exceptions in South
Australia, have been made of the ecology of dry regions. Our data are obviously
incomplete, and must be accepted as inevitably so.

In the present study attention has been almost exclusively focussed on the
climatological aspects of the problem, and we have immediately to decide which
factors of climate are available for correlation. There is no doubt that a full

108 SOME CLIMATOLOGICAL ASPECTS OF ARIDITY,

and detailed examination of climatic conditions at any point or points should
include as many of the so-called climatic factors as possible, viz., precipitation,
temperature, evaporation, humidity, wind, atmospheric pressure, and so on. Such
data are actually available for the capital cities of Australia, as well as for a
very small number of other stations, but from the nature of the case we can
expect only very sparse and most generalized data from the little-inhabited dry
regions of the continent. Indeed, we might say that for the greater portion of
Australia there are available only records of precipitation and temperature, the
latter being given as a twenty-four-hourly mean which is obtained from the
mean of the 9 a.m. and the 3 p.m. readings. In the dry areas the stations are
scattered, while one large region in the central west has no permanent stations.

Maps of relative humidity (Taylor, 1916), evaporation (Hunt, 1914) and
saturation deficit (Prescott, 1931b) have been published. When considering these,
particularly with reference to the climatic ratios described later, the authors
came to the conclusion that in view of the conditions described above, there was
a considerable element of danger in the use of any of them for the purposes of
defining the nature and extent of aridity. As regards the first of these maps,
mean annual relative humidity figures, or even, if such were available, daily mean
figures are of little importance if we do not know the variations which occur
throughout the day, and which have been shown to be very important factors
in the growth of certain plants in arid regions. As regards the map showing
evaporation, there is now a considerable body of opinion opposed to the use of
evaporimeter readings, as the conditions governing the evaporation of water
in an open dish are very different from those governing the evaporation of water
from the soil in the same area. We still have no generally accepted method that
can be applied to continental areas of determining the rate at which water is
evaporated out of the soil, a vitally important matter in regions where duricrust
products are so abundant and where vegetation generally is so dependent
on the presence of water supplies in the soil and subsoil. In the case of the
third of these maps, the concept of saturation deficit is still a comparatively new
one, and as regards Australia the values on which the map was constructed had,
with the exception of a small number of stations, to be calculated from the
mean temperature and relative humidity data. In the first place the objections
previously stated in regard to relative humidity figures hold here too, while in
the second place there is still doubt as to the relations between saturation deficit
and evaporation in arid regions.

It has seemed advisable, then, to work with the first-hand data that are
available and to confine ourselves to precipitation and temperature and their
seasonal incidence. It should be noted that in so doing we are following the lines
laid down by two climatologists who have worked on the continental scale, namely
Koppen (1923) and de Martonne (1927). As is shown later, much of de Martonne’s
work is adopted for the present paper, though it is amplified by considering
monthly conditions and by introducing the concept of the arid period and its
duration.

Index of Aridity.

In an attempt to give a numerical value to the intensity of arid conditions,
several climatic ratios must come under review. Some of these ratios were devised
for quite other purposes than the present one, and are not at all suitable for use
in arid regions. The most important are:

BY J. ANDREWS AND W. H. MAZE. 109

1. Transeau Ratio.—This ratio may be expressed as as the relation of annual
precipitation (P) to annual evaporation (E). It was thought that this would
give an index of effectiveness of precipitation chiefly as regards plant life, but
it is obvious that for States involving areas of continental size the value of the
ratio is diminished by the small number of stations taking evaporation readings.
In Australia there are available only some 23 stations with published evaporation
figures, and these are mainly outside the arid regions. Also, five of them are
situated on the coastline.

2. Thornthwaite Ratio—In a recent study of the climates of North America,
C. W. Thornthwaite (1931) faced the same difficulty, and devised an empirical

12
method whereby the — quotient may be computed from the mean monthly

temperatures and the mean monthly precipitation. The method is as follows:

12
The quotient — is considered the precipitation effectiveness ratio of a single

12?
month, and the = index is the sum of the twelve monthly = ratios.

T in degrees F., and P in inches. To avoid the inconvenience of dealing with
fractions, each quotient has been multiplied by ten, so that the precipitation

12
effectiveness index is ten times the sum of the twelve monthly — ratios. It is
E

12
clear that this — index is not the same as Transeau’s, which is the relation of

annual precipitation to annual evaporation. This equation is checked by reference
to the fairly large number of stations taking evaporation figures in North America,
but at the same time it must be remembered that this apparently precise method
of determination may very possibly have little basis in reality. In arid regions in
particular, there are no means of checking the relations which are assumed
between precipitation, temperature and evaporation. For all essential purposes,
Thornthwaite’s ratio must be regarded as an attempt to carry further the
Transeau ratio, and as a method which, while apparently giving approximately
correct results when applied to North America, could obviously not be extended
in toto to areas in which primary figures are much less satisfactory.

P
3. Meyer Ratio.—The Meyer ratio, —, is the relation between the precipitation

in inches or centimetres, and the aEioepnenie water vapour saturation, deficit,
also measured in inches or centimetres. Even more than the Transeau ratio this
can only be applied to those very rare stations which take measurements of these
factors, and thus can probably be applied to no large continental areas. However
more exact may be the measurement of the saturation deficit as an index of
precipitation efficiency than is the measurement of evaporation, it is at present
impossible to argue with any degree of assurance concerning conditions in arid
regions.

110 SOME CLIMATOLOGICAL ASPECTS OF ARIDITY,

4, In recent studies of the soils of Australia J. A. Prescott (1931la, 1931b)
has used the Meyer ratio and produced maps of atmospheric saturation deficit
for Australia. The saturation deficit has been calculated from the relationship
between the mean annual temperature and relative humidity. The mean
evaporation in inches is said to be related to the saturation deficit in inches of
mercury as E = 263 s.d., this figure having been obtained by the correlation of
23 stations. By means of these figures the Meyer ratio and the Transeau ratio
are considered as being more or less interchangeable. The present authors
consider, however, that a map constructed on this basis and with these data
can only be so highly schematic as to have little value for the purpose of the
delimitation of climatic boundaries. Relative humidity figures themselves are
scarce, and the determination of saturation deficit by means of these can only be
obtained for a number of stations and in certain localities, On the other hand,
it should be pointed out that the precision given to the relation between saturation
deficit and evaporation is in all probability more apparent than real. It is
particularly questionable when applied to tropical and central portions of the
continent, since stations available for the correlation are few in number and
are to be found only in southern Australia.

P
5. Lang Factor—The Lang factor, rae is the quotient of annual precipitation

in millimetres divided by the mean annual temperature in degrees centigrade.
This factor has been criticized, firstly because it assumes a constant ratio between
precipitation and evaporation, secondly because it considers only these two factors,
P and T. As regards the first objection, it is true that this simple relationship
does not obtain over a wide range of climatic conditions, while as regards the
second, the paucity of our information concerning evaporation, saturation deficit
and so on is at least as great as our ignorance of wind and other factors, particu-
larly in arid regions.

6. A further index has been devised by de Martonne, which is in effect an

extension of the Lang ratio. This is to be expressed as , where the precipi-

tation in millimetres is related to temperature in degrees leontionde! The intro-
duction of the constant in effect obviates some of the anomalies resulting from
the use of the Lang factor. It is probable that a more detailed use of de Martonne’s
ratio would help towards the reconsideration of the value of the constant, but
at present, on the basis of physiographic grounds, the constant is accepted as
fixed. Important features concerning this index are as follows:

(a) It makes use of the only reliable climatic data which are available for
that part of the continent. The relation it expresses is simple and yet
fundamental, and may be modified by the introduction of other factors
as our knowledge increases. Empirically, the ratio has been tested for
all the continents of the world, and has been found to agree well with
physiographic indications of climatic regimes.

(6) Significant index numbers.—De Martonne considers the significant values
to be as follows: Indices below 5 characterize true deserts; indices about
10 more or less correspond with prairies; about 30, forest vegetation
tends to predominate, and gains complete control of the soil where the
index exceeds 40, provided the temperatures are not too low. In the
same way no cultivation is possible without irrigation where the index

BY J. ANDREWS AND W. H. MAZE. 111

is below 10. Between 10 and 20 is the domain of dry farming. (Compare
these with Fig. 1.)

(c) While allowance is made in the formula for the varying relations between

effectiveness of precipitation and the temperature, the factor of seasonal
concentration of precipitation does not receive adequate attention if mean
annual conditions alone are considered.

The Intensity of Aridity.

Examination of the intensity of arid conditions is the first step in defining
the nature of aridity. Using de Martonne’s index a series of maps of this nature
were constructed. Fig. 1 shows the annual conditions.

The Map of Annual Conditions.*

Features that may be observed are:

a

In general arrangement and pattern the lines of equal aridity resemble the
arrangement of the annual isohyets, i.e., they are (a) concentric with the
central regions of the continent; and (0) the index numbers increase
towards the coasts, i.e., aridity decreases.

There are some interesting divergences from the above general impression:
Aridity isolines which coincided with isohyets in southern Australia lie
considerably further to the north of the same isohyets in tropical Australia;
so that the introduction of the T factor has enlarged the arid boundary
in tropic areas which were not classed as arid solely by consideration of
the P factor.

There is little differentiation in the central regions. This is mainly due
to the small number of stations maintained therein; the figures available
show variations that suggest fairly complex conditions.

There are two areas of very high values, both situated in the eastern
portion of the continent. The first and more extensive of these is on the
North Queensland coast in the vicinity of Cairns. Here values reaching up
to 109 (Innisfail) may be obtained, these being mainly the consequence of
very high rainfalls experienced along this littoral. The second region is
the Monaro and the Australian Alps, where fairly high rainfalls are
combined with low highland temperatures.

There are some interesting correlations to be made with various other
distributions. The isoline of ten includes practically all the country in
Prescott’s vegetation map marked as desert sandhill and as acacia semi-
desert, and also some of the mallee and sclerophyll woodlands. Rain forest
is found only where the index rises to 40, and where temperatures are
high. In regions of similar index (40) and low temperatures, as in
Monaro, mountain grassland and moor are to be found. The regions
between 10 and 20 which are situated north of the tropic include most
of the savanna country of the continent, and similar regions in southern
Australia are occupied by mallee and by sclerophyll scrub and woodland.
Unfortunately, it is not possible to distinguish accurately de Martonne’s
true desert areas which have an index of less than 5, though their eastern
borders are indicated. ‘There is therefore no differentiation in our map

* This map and the maps of monthly conditions have been drawn with the aid of
numerous rainfall and temperature statistics supplied through the courtesy of the
Commonwealth Meteorologist, Mr. W. S. Watt.

112 SOME CLIMATOLOGICAL ASPECTS OF ARIDITY,

corresponding to the differences between sandhill desert and Prescott’s
acacia semi-desert and shrub steppe. As regards the agricultural distri-
butions, it will be seen that the main wheatlands of the continent lie in the
regions with index 12 to 30, and that there is no agriculture under natural
conditions in regions with index lower than 10. Tropical agriculture is
only to be found in those regions with an index of more than 35.

| HF Ore aT | ea
1s 120 125 SSS 10. 9 140 ON 145 150
rs
|_—-20
{
| ROPIC
pe

|

te) 200 400 NES

JANUARY.

5 120 125 130 ii 140

\ \ oo

Fig. 3.—Index of aridity for January. The lower the index figure the more intense is
the aridity. The isoline of index 1 is significant in distinguishing the arid border.

6. The relations of aridity isolines and the nature of the drainage system
may be summarized as follows: It is not possible to give a numerical
value to the boundary between areic and endoreic areas.* The boundary
between endoreic and exoreic regions, however, may be drawn with
considerable accuracy. This will be found to coincide fairly closely with
the isoline 10, except on the north-west coast from Shark Bay to Port
Hedland, where several occasionally flowing streams such as the Ashburton
and the Gascoyne have their channels emptying to the Indian Ocean.

* These terms were proposed by de Martonne. Regions of areism are those with
no run-off of precipitation; of endoreism, those of interior-basin drainage; of exoreism,
those drained by streams flowing to the oceans.

BY J. ANDREWS AND W. H. MAZE. 113

‘The annual map, then, gives a general impression of the conditions which
agrees well with other observed facts. At this stage it is interesting to compare
the regions indicated as arid in this map with the arid regions shown in Koppen’s
map of the continent (Koppen, 1923). It will be seen from Fig. 2 that Koppen’s
diagram divides the central regions into “desert” (BWh, desert climate, reduced
rainfall less than 25 cm., mean annual temperature greater than 18° C.), and
“steppe” (BShw, steppe climate, reduced rainfall 25 to 50 cm., hot, with mean
annual temperature greater than 18° C., dry season winter; and BSks, steppe
climate, reduced rainfall 25 to 50 cm., winters cold, mean annual temperature less
than 18° C., warmest month greater than 18° C., driest season in summer). The
southern boundary of Koppen’s “steppe” regions coincides fairly exactly with the

200 400 WiLes

MARCH.

130 135

Fig. 4.—Index of aridity for March.

aridity isoline of index 10 (Fig. 1), but in the tropical north the latter is rather
coincident with the “desert” boundary of Koppen. In other words, Koppen sees
steppe conditions extending further north in the tropics. This same tropical
region of the South Kimberleys appears in Prescott’s vegetation map of Australia
(Prescott, 1931) as “savanna woodland” mixed with “savanna and Mitchell grass
downs”. The region is draiied by the Fitzroy and the Ord, which both have well
defined channels. The disharmony results from considering annual summations
alone in a region with very strongly marked seasonal characteristics.

The annual map must necessarily, however, be a generalization, as monthly
conditions vary very considerably in different parts of the continent. It is

114 SOME CLIMATOLOGICAL ASPECTS OF ARIDITY,

essential to a full understanding of the nature of aridity that monthly conditions
should be analysed. A series of monthly maps using the same formula was
constructed and those for January, March, June and October were selected as

typical of the seasonal conditions. The most important features of these may
be summarized as follows:

1. During January (Fig. 3) the most arid portions of the continent are to be
found in the south-west and central southern areas. High indices repre-
senting heavy summer rainfalls can be found through most of northern
Australia, and extending down the east coast. The gradient is very steep
inwards both from the north and east coast as distinguished from the
very gentle gradient of the dry area. In tropical Australia, the isolines

ue i is re ib I

15)

0 200 4

OW iLES

JUNE.

13S

Fig. 5.—Index of aridity for June.

run east and west with only slight irregularities until the east coast is
approached. During this month there is practically no agriculture within
the isoline 1. South-west portions of the continent are under prolonged
summer drought. This applies also in rather less degree to the wheat-
lands of South Australia, north-west Victoria and the plains of New
South Wales.

2. Conditions in March (Fig. 4) show considerable similarity. The isoline 1
has moved westward from the Riverina, but the arid areas are increased
in the centre of the continent, where the autumn rains are very sparse.

BY J. ANDREWS AND W. H. MAZE. 115

The extreme south-west portions of the continent are now receiving their
first winter rains. In the tropical and east coast areas the gradient is
very much less, though almost identical values are received in the Cairns
section as were received in January. As regards agriculture in southern
Australia, the commencement of the wheat sowing season is marked by
the westward retreat of the isoline 1. In south-western Australia, on the
other hand, where rather later sowing rains are desirable, summer droughts
are still in force.

3. The June map (Fig. 5) shows a complete reversal of the January conditions.
The arid portions of the continent are now north and north-west, with the
east coast and the south of Australia receiving their winter rainfalls. The
agricultural lands of the south are now showing indices of over 2, and the

|

1S 120 12s Wem Ve ob 67 150
r es ie
¢ m9 {

20 —~_|

| OCTOBER ;
at ee eee ea 78

Fig. 6.—Index of aridity for October.

wettest portions are the southern littorals. It is worth noting that this
arid area in the winter month is considerably more extensive than is the
arid area shown in the January map.

4. For October (Fig. 6) the chart shows extensive location of arid conditions
as far as northern and central Australia is concerned. The agricultural
lands of the east coast and the Murray-Darling basin alone in the east
show higher indices than 1, while in the whole of the western half of
the continent only Swanland has similar values. This month is agricul-
turally important in the determination of the ultimate yield of the wheat

116 SOME CLIMATOLOGICAL ASPECTS OF ARIDITY,

crops, and climatic conditions at this period are extremely important in
influencing the extension of wheat crop areas. On the extreme north
coast, centring round Darwin, the “Wet” is beginning to appear, and
Darwin shows a value of 1:3.

TABLE I.—Indices for Important Stations.

January. March.
12) T I P dE If
Sydney Se ne A ae 91-4 21:9 2-90 126-0 20-7 4-10
Melbourne... a3 ae ce 47-8 19-6 1-61 Bisel 18-1 2-03
Brisbane Ss as wi zfs 165-0 Zor 4-70 148-0 23-5 4-40
Perth .. ies ae ee ao 8-6 23-2 0-26 19-8 21-8 0-62
Adelaide ae bcs ee is 18-0 23-2 0:54 25-6 21-0 0-86
Darwin Es ee 3 ae 398-8 28:8 1:01 253-0 28-9 6-50
Broome Ale ibs Fins ‘a 160°6 29-7 4-05 89-4 29-6 Ze
Alice Springs re Eye Bio 43-4 28-4 1:14 31-2 24-8 0-90
Camooweal .. 56 pre oe 98-6 30°4 2:45 53°6 28-1 1-40
Broken Hill .. ie ats a iyo 25-6 0-48 16-0 22-1 0:50
Wiluna 4 Me a ed 34:8 29-8 0:87 32-0 26-6 0-87
Hall’s Creek .. Be: as ae 147-0 30°3 38°64 (HOV 28-2 2-02
Albury nie fe oe ae 38-6 24:3 1-13 49-9 20-6 1-63
Tarcoola is 55 os ss 8-4 24-7 0-24 12-2 23-3 0:36
Daly Waters 2: ae ae 162-6 30°5 4-00 120-9 28-7 3:12

TABLE I.--—-Indices for Important Stations.—Continued.

June. October. Year.
12 a I P Av I 1 T I
Sydney ie go || IeBeal 12-6 5:40 73°9 17°6 2-67 1,206 17-3 44-4
Melbourne .. ts 52:3 10-2 2-60 66-0 14:3 2-70 648 14:7 26-1
Brisbane .. ore 71-6 iso? 2-78 64-3 21-0 2:07 1,157 20-5 38-0
Perth ae sa- |) aizkshe'5 1183 ¢/ 7-50 54-7 16-0 2-10 883 17-9 31-6
Adelaide .. Bs 79-2 11:9 3-60 43°7 16-7 1-64 536 17-2 19-6
Darwin on aes S20 25-9 0:08 51:8 29-6 1:30 1,531 28-2 40-0
Broome ate ss 24-4 21-8 0:77 0-7 hie: 0-02 590 26-5 16-2
Alice Springs be 14-7 12-4 0-65 18-8 23-0 0-57 272 20-9 8-8
Camooweal oe 9-4 17-9 0-32 13-9 27-3 0:37 385 24-9 alow
Broken Hill FES 30-2 10-7 1-45 20-3 18-7 0:71 244 18-1 8-6
Wiluna ats Me 25-4 12-9 aaa la 6°3 21-4 0-20 250 21-4 7:9
Hall’s Creek Re 4-5 18-7 1-60 Io) 28-6 0:39 524 25°5 14-8
Albury Ae ae 85-5 8-9 4°51 65:8 157, 2-60 692 16-0 26-6
Tarcoola .. ate VA hi 10:9 1-60 17°8 18-6 0-62 177 18-4 6-2
Daly Waters 5 6-3 21-3 0-20 21:3 30-1 0°53 670 26:8 18-2

5. The arid border is a zone which fluctuates from month to month. In bad
seasons it may extend into whole divisions of any of the States and
signalize crop failure. In good seasons the wheatlands flourish, and far
into the interior grass and water are abundant. The defining of this zone
will be partly a matter for field work, but it can be partly defined by our
existing knowledge. From careful examination of the statistics, supple-

BY J. ANDREWS AND W. H. MAZE. 117

mented by our knowledge of the behaviour of some crops under certain
climatic conditions, a tentative selection of the index 1 was made as the
arid boundary. Knowledge of the distribution of wheat cultivation and
the growth of wheat under certain climatic conditions is very useful, for
wheat is the most widely-spread and one of the most uniformly cultivated
of our crops. (See Table II for representative occurrences of the index 1.)

TABLE II.—Svme Representative Stations Showing P and 7' Values, which give a Monthly Index of 1.

Month
in which
Station. Lat. 8. Index. that T in that P in that
Index Month. Month.
Occurs.
F° (Oy in mm
N.S.W.—
Bourke a oat 30°5 1-00 May 59-8 15°5 1-01 25:6
Pe 1-05 Feb. 83-0 28:3 1-59 40-4
if Be € 0:92 Dec. 82°3 28-0 1-37 34-8
Broken Hili ste 31°58 0°99 Aug. 52-9 11:6 0-84 21:3
“A ays 1:02 May 56:6 118307/ 0:97 24-6
Forbes ae Ne 33°25 1-10 Feb. 77°6 25:4 1-54 39-1
EV averse 8 Ns 34°35 1-00 Apr. 62-9 ilyfoul 1-08 27-4
Quambone .. 53 30°31 0:99 Jan. 80°7 27-0 1-45 36°8
Walgett Sis ¥: 30-1 1-00 Oct. 69-4 20°7 1LSPAl 30-7
Nyngan se ae 31-34 0-96 Sep. 58-6 14:8 0-94 23-9
Victoria—
Mildura Se oe 34-13 0-94 Oct. 63-4 17-4 1-02 25-9
Charlton es Sie 36°16 0-98 Mar. 66°5 19-2 ilaly/ 29-7
Bendigo an ae 36:46 0:96 Feb. 71-3 21-8 Toph 30°7
South Australia—
‘Adelaide te Se 34:57 0-98 Nov. 67-0 19-4 1-14 28-9
Port Pirie .. Seg 33-1 0:96 Apr. 67°2 19-5 itp 28-4
Queensland—
Thursday Island .. 10°33 0-96 Nov. 82°7 28-2 1-45 36°8
Townsville .. Be 19-10 0:97 May 15°3 24-1 1-30 33-0
Windorah .. << AHORA 0:97 Dec. 85-4 29-6 1:51 38-3
Charleville .. a 26°55 0:97 Oct. 72-4 22-4 1-24 BLOG)
Bollon re es 28-0 0-99 Oct. 70-7 PALO} 1-23 31-2
Western Australia-—
York .. ae a 31-53 1-00 Oct. 59-8 L515 1-01 25-6
Kalgoorlie .. se 30-45 0:97 =! July 52-4 11:3 0-82 20-8
Hall’s Creek. . ae 18-16 0-95 Nov. 87-1 BiNo7/ 52 38-6
~ Derby ae Bs 17-19 1-05 Apr. 83°5 28-6 1-60 40-6
Marble Bar a 21:6 0-99 June 67-7 19-8 1-16 29-4
Central Australia—
Alice Springs a5 23°42 1:02 Feb. 82-2 27-9 1-52 38-6

The Duration of the Arid Period.

Having made a selection of the aridity index 1 as a significant indication of
the extension of arid conditions, Figure 7 was plotted so as to show the number
of months through the year which in average years would be classed as arid.
This brings us to the consideration of the duration of the arid period and its
position in the year, a point which we regard as of primary importance. The
general features of the map may be summarized as follows:

1. There is an area in the central southern portions of the continent which
has practically continuous arid conditions. This extends from the south-

118

SOME CLIMATOLOGICAL ASPECTS OF ARIDITY,

western portions of Queensland, through the northern portion of South
Australia to the Western Australian boundary. Its westward extension
from this line cannot be precisely determined because of the absence of
meteorological stations, but it is probable that the central portions of
Western Australia (i.e., a high proportion of the area containing sand
ridges) come within its boundary.

i 125 ITA ney

WS 120 125 130 135 @ 150

\ \ | | | l

Fig. 7.—The mean duration of the arid period, in months. Regions with an index of
12 have practically permanent aridity; those with an index of 0 are rarely arid for any
months. The index of 8 is suggested as significant in the definition of the arid border.

2. There is an extremely steep gradient in eastern Australia, and particularly

in south Queensland, between the extremely arid and the humid areas. It
is to be interpreted in the first place as indicating a marked and sudden
change in arid conditions which should be reflected to some comparable
extent in landscape and vegetation distributions. In the second place it
will be obvious that in these regions deviations from the average conditions
will produce very marked effects due to the close propinquity of arid and
humid regimes.

The regions in which arid conditions do not exist in average years are
limited to areas in which precipitation is fairly evenly distributed through
the year. These regions are confined to eastern Australia. The most
favourably placed areas are indicated as being south-eastern Queensland,
and New South Wales and Victoria to the east of the highlands and

o 20 —_|

BY J. ANDREWS AND W. H. MAZE. 119

western slopes. A small isolated area of similar characteristics is to be
found on the north Queensland coast in the vicinity of Cairns.
Interesting conditions are to be observed in south-western Australia. Here
there are at least four months of arid conditions. These dry periods,
which occur mainly in summer and spring, inhibit growth for the time
being, but have not prevented the development of extensive forests and
woodlands. ‘

Tropical Australia shows, with the exception of the Queensland coast,
comparatively little range in arid conditions. The dry period ranges in
length from five to ten months, but the greater portion is less than eight
months.

As an indication of the arid border, considerable reliance is to be placed
upon the isoline of 8. This coincides well with the extent of desert and
semi-desert vegetation associations; includes the area having endoreic
drainage; and there is included within it no lands in which agriculture
has proved successful, though it comes close to the dry border of the
important wheatlands.

As compared with the map showing the intensity of the annual conditions
(Fig. 1), this duration map provides a much more satisfactory summary.
Since aridity means the temporary suspension of many biological and geo-
morphological processes of fluvial regions, then an important question is
the average length of the period which elapses before these are resumed.
Thus from the biogeographical viewpoint, the most important aspect of
arid climates is that the arid period is relatively very long and not that it
is relatively dry. Other climates (such as tropical climates which have
a dry season, or that group of climates resembling more or less closely the
Mediterranean type) also have an arid period which, however, in average
years is not long enough to result in the establishment of true arid plant
and landscape types.

It follows from what we have said that there is a critical length of the
arid period. Regions where a dry period longer than this critical length
recurs frequently are desert-like, though an occasional retreat of the arid
border means a brief time of luxuriantly growing annuals and flowing
stream channels. On the other hand, regions which usually lie on the
wetter side of the arid border sometimes experience years when the arid
period stretches beyond the critical stage, and drought takes possession of
the land. In the central west of New South Wales one may see areas of
dead pine which were killed in the 1902 drought. Drought is, after all, a
relative term, and is applied to dry periods which last longer than is normal.
The final point to be made here deals with the frequency of the occur-
rence of the critical length. Every climatic type represented in Australia
has an arid period of greater or less length. What marks out large areas
of the interior from the other regions is the greater intensity, duration
and frequency of the arid period. The question of frequency will lead
us to the consideration of actual instead of average years, but we know
that in these regions it is frequent enough to result in the establishment
of certain recognizable types of landscape and vegetation. One of the major
problems that confronts us is concerned with the relation of the intensity
and frequency of the arid period and the cumulative effect on the natural
landscape of very frequent arid periods of more than critical length.

120 SOME CLIMATOLOGICAL ASPECTS OF ARIDITY.

Conclusion.

The present-day climatic factor is the most important of the immediate causes
of aridity. The dominant climatic character in arid regions is the length of the
period in which most geomorphological and biological processes of fluvial regions
are suspended or greatly curtailed because of the absence of an effective precipita-
tion. Definite limiting values are suggested for the efficiency of precipitation
(in monthly means) and for the critical length of the arid period. The authors
suggest, as a basis for discussion, the definition that regions of aridity are regions
of markedly intermittent and strongly contrasted geomorphological and biological
processes which are controlled in their occurrence by the length of the period of
insufficient precipitation; these arid periods are of such length and recur so
frequently that the processes associated with them are dominant in determining
the cultural and natural landscapes.

References.

Hunt, H. A., 1914.—Climate and Meteorology of Australia. Comm. Bur. Meteor., Melb.,
Bull Noe 931914 ps ib?

DE MArTONNE, EMoM., 1927.—Regions of Interior-basin Drainage. Geog. Rev., xvii, July,
UBYATS 105) Bete

JEFFERSON, MArK, 1916.—Aridity and Humidity Map of the United States. Geog. Rev.,
i, 1916, p: 203.

KOPpPEN, W., 1923.—Die Klimate der Erde. Berlin, 1923.

Prescott, J. A., 193la.—The Soils of Australia in Relation to Vegetation and Climate.
Counc. Sci. Indust. Research, Com. of Aust., Bull. 52.

,1931b6.—Atmospheric Saturation Deficit in Australia. Trans. Roy. Soc. Sth.
ANSE, Iva 93d:

RUSSELL, R. J., 1931.—Dry Climates of the United States. Univ. Calif. Publ. Geog., 5,
WO; il; joy i, ;

TAYLOR, GRIFFITH, 1916.—Control of Settlement by Humidity and Temperature. Comm.
Bur. Meteor., Melb., Bull. 14, 1916, figs. 5, 6.

, 1920.—Australian Meteorology. Oxford Univ. Press, 1920.

THORNTHWAITE, C. W., 1931.—The Climates of North America according to a New

Classification. Geog. Rev., 21, Oct., 1931, p. 633.

SEASONAL INCIDENCE AND CONCENTRATION OF RAINFALL IN
AUSTRALIA.

By Joun ANpreEws, B.A., Demonstrator in Geography, and W. H. Mazz,
Caird Scholar in Geography, University of Sydney.
(Five Text-figures. )
[Read 26th April, 1933.

A series of maps showing the proportion of the annual rainfall oceurring in
each season is useful for many purposes dealing with climatological, pedological
and vegetational distributions. A complete series does not seem to have been
published before in Australia. In each case the maps given here (Figs. 1-4)
show the percentage of the yearly rainfall which falls during the season in
question.

A further map (Fig. 5) which has been found useful is that showing the
degree of concentration of rainfall in the wettest season. This is constructed
from the values for each station obtained by taking the difference between the
highest and the lowest seasonal percentage amounts. The table shows these
values for representative stations.

200 400 ices

Exar : p a SU MMER. 140
Me Tbe ee

Fig. 1.—Summer rainfall, showing the percentage of the mean annual total that falls in
December, January and February.

M

3122 SEASONAL INCIDENCE OF RAINFALL IN AUSTRALIA,

é ws
a o—
|
29 do
|
40
rROPIC TROPIC
30. |
| }
50;
| A
aie an
| |
! fi
© 200 ACS yin ex | VA
ee 35 i sl
| AUTUMN. . i?
ws 206 25 130 135 140 150
Peas or LP ee ney aS oe

Fig. 2.—Autumn rainfall, showing the percentage of the mean annual total that falls
in March, April and May.

_ WINTER.

| 130 135 140

| 30

Fig. 3.—Winter rainfall, showing the percentage of the mean annual total that falls in
June, July and August.

BY J. ANDREWS AND W. H. MAZE. 123

WS

20

MILES

yc
z SPRING . i
ts 12e i 130 140 Siete 7

Fig. 4.—Spring rainfall, showing the percentage of the mean annual total that falls in
September, October and November.

'SEASONA L CONCENTRATION oh oes |

\ | | | | 19 | 2
Fig. 5.—The seasonal concentration of rainfall. The index figure is the difference
between the highest and the lowest seasonal percentage amounts. Seasonal maximum
regions are also shown (bounded by dotted lines).

124 SEASONAL INCIDENCE OF RAINFALI. IN AUSTRALIA.

Seasonal Rainfall at Representative Stations.

| j

Summer. Autumn. Winter. | Spring. Concentra-

Station. % %, % | % | tion Index.
Thursday Island ay Lys a 60-6 35-1 1:6 Bol 59-0
Normanton .. fs as a 72-0 20°3 1:5 6-2 70°5
Moree pi ae ee a 32-4 24-3 20-6 Hi 11:8
Bathurst Bie ae an Ae 29-0 bbe PIO 7 25-6 5-3
Salevia of ay RC aft 24-1 24-6 Pala? 29-6 7-9
Sydney sa au 22:5 33-0 26-6 17:9 15-1
Cape Leeuwin we ae ae 6-1 24-0 50:2 | 19-7 44-1]
Laverton A ae re ae OA /O5) 37-1 ORL | B3}0%/ 15-4
Onslow te ae as a: 23:8 41-8 33-4 1-0 40-8
Alice Springs es ats ee 44-5 23-9 i24oa | 19°5 32-4

(Seasonal figures are percentages of the mean annual total.)

In the map (Fig. 5) the index figure may be regarded as an index of
concentration, the name of the season in which maximum concentration occurs
being given also. Thus most of New South Wales has a low degree of concentra-
tion, there being a difference of only 10 (more or less) between the highest and
the lowest seasonal percentage figures. The shores of the Gulf of Carpentaria,
on the other hand, have a highly concentrated rainfall, since the season of
heaviest fall (Summer) has at least 70% more of the year’s total than has the
driest season. It follows in this case of course that the other seasons must
also have small percentage figures. The actual figures for Normanton are given
above.

The boundaries showing the season of greatest fall differ in some particulars
from other published maps.

THE PETROLOGY OF THE HARTLEY DISTRICT. II.
Ture METAMORPHOSED GABBROS AND ASSOCIATED Hysrip AND CONTAMINATED Rocks.

By Germaine A. Joprin, B.Se., Curator of the Geology Department Museum,
University of Sydney.*
(Plates v—vi; four Text-figures. )
[Read 31st May, 1933.]
Introduction,

In the following pages an account is given of certain rocks of mixed origin
that have been produced as a result of (1) interaction between a solid gabbro
and a quartz-mica-diorite magma, (2) contamination of the gabbro by sediments.
The thermal metamorphism of the gabbro, due to the emplacement of the more
acid rock. is also considered.

The writer has previously (1931) described a calcic plutonic complex ranging
from acid granites through an intermediate and basic series down to the ultra-
basic type—hornblendite. Chemical analyses were made to show the consanguinity
of the types, and although it was realized that the gabbros had suffered contact
metamorphism, the present more detailed study of these phenomena has revealed
the fact that the rock analysed for a pyroxene-gabbro (Joplin, 1931, p. 43) is a
hybrid formed by the interaction between the solid gabbro and quartz-mica-
diorite magma. Further studies, however, the examination of ninety-one micro-
sections, and two more analyses have shown the two intrusions to be comagmatic
and the rock previously analysed to be but slightly acidified. Furthermore, it
can now be shown that the hornblendites are hybrids and of the nature of basic
segregations.

The mass under examination has been referred to previously as the Cox’s
River Intrusion, and outcrops on Cox’s River about 3 miles below the Glenroy
Bridge on the Jenolan Road. A small portion of the mass outcrops within the
Parish of Hartley, County of Cook, but the greater part of it lies to the west of
the river in the Parish of Lowther, County of Westmoreland.

Field Relations.

The Cox’s River Intrusion, and the Moyne Farm Intrusion, a composite
diorite stock 14 miles to the east (Joplin, 1931), are injected into the trough of a
syncline trending approximately east and west and pitching to the east. This is
occupied by Upper Devonian sediments and lavas. Granite outcrops to the north
and south of the syncline and its junctions are roughly concordant with the
strike of the sediments. Both to the north and to the south the broad syncline is
turned over abruptly into a sharp anticline against the margin of the granite.
This possibly may be due to a drag effect of the invading granite.

The Cox’s River Intrusion occupies an area of about 900 acres, but a fairly
large outcrop of quartz-mica-diorite in Por. 52, Parish of Lowther, suggests that
the acid phase is more extensive than at present appears to be the case. The
centre of the main mass is situated in Por. 137, Parish of Lowther, and the rocks
at the junction of the creeks at this point appear to be normal hornblende-

* This work was commenced when the writer held a Sydney University Science
Research Scholarship in Geology and the Deas-Thomson Scholarship for Mineralogy.
A

126 PETROLOGY OF THE HARTLEY DISTRICT, ii,

gabbros that have suffered a slight deuteric alteration. Reference to Plate v
shows that passing out radially in any direction a recrystallization becomes
apparent. A partial granoblastic structure is developed, and monoclinic and
rhombic pyroxenes are the only ferromagnesian minerals. Further out still
hornblende again makes its appearance and a fabric resembling the primary
ophitic gradually comes into evidence. Isolated outcrops of diorite-gabbro and
hornblende-gabbro occur among the hornblende-pyroxene-gabbros, and scattered
through these latter and also through the recrystallized pyroxene-gabbros are
rocks that have suffered hydrothermal alteration, the pyroxene having been
changed into pale uralitic material. A mass of quartz-mica-diorite, varying in
width from 10 chains to half a mile, completely encompasses this gabbro complex,
and it has been pointed out already that there is reason to believe that these
rocks are even more widespread.

Form of the Quarte-mica-diorite Intrusion—Although the quartz-mica-diorite
encircles the gabbro, there is some evidence to show that the engulfment was not
of a normal type. It seems evident that the junction between the diorite and
the gabbro is convex upwards and that the thickness of diorite on top of the
gabbro was not very appreciable.

Plate v shows that near the head of Marriott’s Creek there is a mass of
quartzite (15 x 10 chains) resting partly on the surface of the gabbro, and
about 10 chains to the west of this a small patch of highly altered porphyrite
occurs. As porphyrites are found interbedded with the sediments elsewhere in
the district, it is probable that this, together with the quartzite, represents a
portion of the roof of the intrusion. This would indicate that either the quartz-
mica-diorites did not cover the gabbro, or that the mass is portion of a “roof-
pendant” (Daly, 1906) protruding down below their level into the underlying
gabbro. A large mass of sediments occurs near the margin of the granite about
a mile further up the river, and a small patch among the diorites on Moyne
Farm, and small outcrops of granite and diorite, at short distances from the
main intrusions, are not infrequent. These occurrences suggest that the sedi-
mentary cover of the bathylith has not yet been completely removed, and
that no very deep denudation has taken place. This, together with the fact that
the gabbros often occupy the higher levels of the Cox’s River Intrusion, inhibits
the possibility of a very deep protuberance of sediment, and points either to a
very thin cover of diorite or to no such cover at all.

Reference to Plate v shows that, although the quartz-mica-diorite is of
fairly constant width, that of the “reaction-ring” of hornblende-pyroxene-gabbro
is rather irregular; and, moreover, that this irregularity appears to bear no
relation to the present width of the diorite. These facts seem to point to a
covering of diorite which was apparently of no very great thickness. An
inwardly curving junction between the two igneous rocks is also indicated.

The form of the intrusion is possibly that of a hollow cylinder with a dome-
like roof and a ring-like outcrop. The annular width is fairly constant and
approximately 20 chains, whilst the diameter of the enclosed gabbro mass is of
the order of 60 chains. There appears to be only one radial apophysis into the
gabbro. This annular form may be accidental, though the intrusion appears to
have some affinities to the ring-dykes so common in certain parts of Scotland
(Bailey et al., 1924; Richey and Thomas, 1930; Richey, 1931).

A wide valley occupies the centre of the intrusion and this is surrounded by
steep hills. The physiography is certainly peculiar and is suggestive of down-

BY GERMAINE A. JOPLIN. 127

faulting, especially in view of the ring-like structure of the intrusion. There is
no evidence of faulting, however, and although the structure of the intrusion
is significant, it is impossible to admit a large sunken area.

PETROGRAPHY.
i. Deuterically altered hornblende-gabbros.

The hornblende-gabbros occupy a small area in the centre of the intrusion
on the property of Mr. Chris Commens, Por. 137, Parish of Lowther, and show
little or no evidence of having suffered contact metamorphism. They are heavy
dark rocks with a variable grainsize, and in the handspecimen are seen to
consist of plagioclase, hornblende and iron ore.

Under the microscope they are ophitic to subophitic, and the grainsize is
usually medium (2 mm.—3 mm.). The constituent minerals are plagioclase, horn-
blende, augite, hypersthene, iron ores, apatite and sometimes a little biotite and
quartz. Epidote, chlorite, kaolin, saussurite, iddingsite, (?) lawsonite and a
zeolite are present as products of deuteric action.

The plagioclase occurs in idiomorphic to subidiomorphic crystals with a
stout columnar or somewhat tabular habit, and is partly wrapped round by,
or included in the femic minerals. The composition is labradorite (Ab,,An,,),
but occasionally a slight zoning is present. Twinning is well developed after the
aibite law, and sometimes pericline twinning is present. A few small indeter-
minate inclusions occur, and the felspar shows a good deal of alteration into
epidote, saussurite and kaolin.

The hornblende forms large subidiomorphic columns or irregular sheets, and
often encloses cores of pyroxene about which it is forming a reaction-rim (Bowen,
1922a). Simple pinacoidal twinning is fairly common, and small inclusions of
iron ore abundant. The colour is brownish-green and the absorption Y> or = Z>X.

Augite sometimes forms subidiomorphic columnar crystals, but is more often
rounded and corroded and surrounded by a reaction-rim of hornblende (Plate vi,
fig. 1). Magnetite grains are usually present as inclusions, and in those rocks
that show slight evidence of recrystallization, the augite has developed schiller
inclusions in restricted patches. In the opinion of Judd (1890) and H. H. Thomas
(1930, p. 239) local schillerization is of a secondary origin and may evidence
thermal metamorphism. Harker (1904, p. 109), however, considers that “the
capricious distribution does not seem inconsistent with a primary origin”. Ina
section showing slight evidence of thermal metamorphism (Text-fig. 1) one set
of schiller inclusions subtends an angle of 84° with the prismatic cleavage of the
pyroxene, and the second set subtends an angle of 137° with the first. In the
same microsection small granules of recrystallized pyroxene occur in the brown
hornblende which is forming a wide reaction-rim about a large columnar section
of original augite (Text-fig. 1). Besides the “reaction” change to primary horn-
blende, the augite often shows evidence of having suffered deuteric alteration,
and has been partly converted into uralite.

Iron ores, apparently consisting of both magnetite and ilmenite, occur in
two generations. The first generation is represented by small inclusions in all
the other minerals; the second by fairly large irregular grains moulding both
felspars and ferromagnesian minerals. Small flakes of biotite are intimately
associated with the iron ores in a few specimens, and in these tiny interstitial
grains of quartz are also present. As the rocks containing biotite and quartz
also show slight evidence of thermal metamorphism, it was at first believed that
the biotite was produced as a result of recrystallization as observed by Dr. Tilley

128 PETROLOGY OF THE HARTLEY DISTRICT, 1i,

(1924) in the contact altered epidiorites at Comrie. As no biotite occurs in the
more completely altered types, to be described below, it is concluded that these
rocks represent slightly more acid differentiates of the hornblende-gabbro.

Text-fig. 1. Text-fig. 2.
- 1.—A, pyroxene; B, local schillerization; C, reaction-rim of brown
hornblende; D, plagioclase; HE, recrystallized pyroxene.
Text-fig. 2.—A, pyroxene-rich zone; B, iron ore-rich zone; C, plagioclase-rich
zone; D, fine granular aggregate; H, coarse granular aggregate.

=)
g
wn
i
=)
gg

(?)Lawsonite (Ransome, 1904; Stillwell, 1923; Joplin, 1931) sometimes forms
lenses in the biotite.

Hypersthene is developed in rounded columnar crystals in a few specimens,
and may represent a product of metamorphism, since it occurs in those rocks
exhibiting incipient recrystallization. Occasionally iddingsite is associated with
the hypersthene.

A zeolite is developed in a few of the hornblende-gabbros. It consists of
masses of either unoriented or radially arranged fibres which occupy interstitial
spaces or veins in the rock. Small veins of a secondary amphibole sometimes
occur, and in these the amphibole fibres are arranged at right angles to the
walls of the vein.

li. Recrystallized pyroxene-gabbros.

The pyroxene-gabbros surround the hornblende-gabbros and, indeed, some
of them cannot be distinguished from the hornblende varieties in the hand-
specimen. They vary from fine-grained types to fairly coarse rocks which can
be seen to consist of plagioclase, pyroxene and iron ores.

Under the microscope the pyroxene-gabbros usually show a blastosubophitic
structure, but a granoblastic structure is often locally developed (Plate vi, fig. 2).
The grainsize is mainly even and medium, and the rocks may be divided roughly
into two types—those in which there is but slight or no apparent recrystalliza-
tion of the felspar, and those in which the felspar is partly granular. The field-
relations of these are quite irregular. The constituent minerals are plagioclase,
hypersthene, augite, iron ores, uralite, a little apatite, a trace of brown horn-
blende and sometimes iddingsite and a talcose mineral.

In those rocks in which only slight recrystallization of the felspar has taken
place, the plagioclase forms columnar crystals about 1 mm. in length. If any

BY GERMAINE A. JOPLIN. 129

granular felspar be present in this type, it forms small equidimensional grains
of about 0:05 mm. In the rocks containing partly recrystallized felspar it forms
either stout columnar or tabular crystals that are indented by the small granular
felspars and pyroxenes surrounding them. In these types the proportion of
granular to columnar felspar is greater, and in a few cases the larger plagioclase
erystals have become so indented by the surrounding granules that the rock
appears to be quite granular. In a few slides there is a slight indication of
bending in the longer columnar plagioclase crystals. The plagioclase is labradorite,
but the composition varies in the different specimens from Ab,,An,, to Ab,,ANg¢,
and zoning is usually absent. In a few specimens, however, inverted zoning is
present. This structure has been noted by Dr. W. R. Browne (1927) in meta-
morphosed dolerites from Broken Hill. The most remarkable feature of the
plagioclase is a peculiar clearing and the presence of groups of minute inclusions.
These inclusions are crowded in the centre of the felspar or arranged in zones
(Plate vi, figs. 2, 6). They usually appear as small, dark brown dots, and in a
few cases tiny granules of pyroxene have been recognized. This clearing and the
formation of minute inclusions has been noted by A. Harker (1904), C. E. Tilley
(1921) and W. R. Browne (1927). Dr. Browne has shown that many of the
granules in the Broken Hill rocks consist of pyroxenes and brown hornblende,
and Dr. Tilley has suggested a similar identification. The completely recrystal-
lized felspars are quite limpid and free from inclusions. In a few instances
the centre of a plagioclase crystal is occupied by a pale brown, anisotropic mineral,
which suggests an incipient alteration into brown hornblende.

In some of the specimens augite is present both as idioblastic, columnar
crystals, which usually show sieve-structure. and as small rounded columnar
crystals or granules crowded between the felspars and indicating the original
ophitic fabric of the rock. The fabric of the rock is thus an intergranular one
produced by metamorphism and may perhaps be termed intergranoblastic. In
many specimens the rounded columnar and granular pyroxene is the only type
present. In some cases the larger crystals have become granulated, though the
original shape of the crystal is preserved. Schiller inclusions are fairly common,
and the crystals exhibiting sieve-structure are crowded with small rounded
inclusions of plagioclase, iron ores and (?)quartz. An intergrowth between
the pyroxenes is quite frequent, and twinning is often developed. Many of the
larger augite crystals are flecked with brown hornblende, suggesting a state of
incomplete equilibrium. In one slide a taleose material has been noticed filling
cracks in the pyroxene. This is possibly due to a later period of hydrothermal
metamorphism, which is also evidenced by the development of a secondary
amphibole in the pyroxenes along cracks in some of the rocks.

Hypersthene shows exactly the same features as the augite, from which it
may be distinguished by its lower double refraction, straight extinction and
pleochroism. The pleochroism is comparatively strong for this mineral, which
suggests a fairly high percentage of iron. This high iron content is further
indicated by the frequent development of limonite along cracks in the hypersthene.
in a few specimens iddingsite is associated with the rhombic pyroxene.

Iron ores are extremely fresh, but occasionally a little granular sphene is
associated and it is probable that both magnetite and ilmenite are present. The
iron ores occur as small inclusions in the other minerals, or as very irregular
grains, up to 1 mm. across, moulding the felspars. These latter are in close
association with the granular pyroxenes, which are crowded between the felspars,

130 PETROLOGY OF THE HARTLEY DISTRICT, ii,

and may represent a recrystallized original iron ore and/or a recrystallization
product of the original ferromagnesian mineral that moulded the felspars.

In most of these rocks apatite is present only in small amount and may
sometimes be absent. When present it forms fairly large, stout columnar crystals,
and is moulded by the magnetite and pyroxenes and occasionally by the plagioclase.

In a few of the pyroxene-gabbros, as well as in some of the recrystallized
types that have suffered reaction and/or hydrothermal metamorphism, ellipsoidal
masses of granular augite, hypersthene, plagioclase and iron ores occur. These
may be up to 10 mm. in length and have a peculiar zoned arrangement of the
constituent minerals (Text-fig. 2). Though not always perfectly zoned, the
following arrangement may usually be made out. In the centre there is a core
of poikilitic plagioclase and moderately large pyroxene and magnetite granules,
or a fairly coarse granular mass of these three minerals. This is followed by a
wide rim consisting of an extremely fine granular mass of the same three
minerals and then by a magnetite-pyroxene-rich zone which is narrower and a
good deal coarser in grain. Outside this again comes a felspar-rich layer, then
a zone very rich in iron ores, and finally one consisting exclusively of pyroxenes.
The wide fine-grained zone, and the coarser-grained central zone show some
variations. In one of the ellipsoids a cruciform arrangement has been noticed.
The coarser area forms a cross which divides the finer outer zone into four
sectors. In another the coarse zone is very wide in comparison to the finer, and
in others again it is quite eccentrically placed. Some sections of these bodies
show a remarkable parallelism of the longer axes of the granules. In the hand-
specimens the ellipsoids are not readily distinguishable and appear as small,
dark, stony masses.

In column I below an analysis is given of a fairly normal type of the rock,
and in column II the results of an analysis of a slightly leucocratic type is shown.

il, Te IDOE

SiO, 44-52 44-79 44-40
Al.O3 21-32 19-56 20°55
Fe.0O; 5:08 CiO15 | G57 Quartz 2-04 0-96
FeO 7:19 7:79 9-26 Orthoclase 0-83 0-56 somal
MgO Be as 6-41 6-16 BSOPAl Albite won| L048 9-96 9-43
CaO aM so) || toe 11-81 11-50 Anorthite S226 47°82 50-32
Na,O 1:25 L320) 1:14 Diopside a 7-42 8-10 5:50
K.O 0-15 0-06 0-19. Hypersthene .. | 18-28 19-14 21-90
H,O+ 0:37 0-64 1-00 Olivine 1:21 — —
H,O — 0-06 0-10 — Magnetite Waa? 8-82 9-51
TiO, 5'6 at 1-04 1-14 — Ilmenite Be 1-9 2-13
P,0; Me aa abs 0-18 — Apatite .. AP — 0-34 =
MnO Se ve 0-09 OMS —
co, 35 se tr tr os

Motaliwoa. .. | 99:92 99-60 99-82

Specific Gravity 3°050 3055 3°035

I.—Recrystallized Pyroxene-gabbro (leucocratic phase). South end of Por. 239,
Par. Lowther, Little Hartley (Corsase, near Kedebekase and Hessose, II(III), 5,
(4)5, 3”). Anal. G. A. Joplin.

IIl.—Recrystallized Pyroxene-gabbro (normal type). North end of Por. 32, Parish
of Lowther, Little Hartley (Kedebekase, near Corsase and Hessose, (II)III, 5, (4)5, 5).
Anal. G. A. Joplin.

IIlI.— Segregation in Norite. The Bluff, Otago, N.Z. (Corsase, II(III), 5, “5, 0).
Anal) i J. Wild. Trans: N.Z. inst. xliv,, 191d) Giga). pi s2b.) Tne aWedl ap. coo 2, NOs 0.

BY GERMAINE A. JOPLIN. 131

These “analyses, particularly the one given in column II, show that the chief
characteristics of the rocks are their high alumina and iron percentages and
low potash. The chemical composition of the gabbro would suggest that it had
affinities with the eucrites, and this is confirmed to some extent by the presence
of very basic labradorite. According to Holmes (1920) a true eucrite contains
bytownite-anorthite and pyroxene.

iii. ‘Reaction’’-gabbros or hornblende-pyroxene-gabbros.

Reference to Plate v shows that the hornblende-pyroxene-gabbros form an
outer border or reaction-ring about the recrystallized pyroxene-gabbros, and the
following petrographical descriptions will serve to show that this group has
suffered recrystallization, as well as reaction with the quartz-mica-diorite magma.

The hornblende-pyroxene-gabbros vary a great deal in the handspecimen.
They are mostly dark, heavy, fairly coarsely crystalline rocks, often showing
large “shimmer” plates of hornblende, crowded with inclusions, and giving both
a porphyritic and poikilitic texture to the rock. Under the microscope the
structure is partly granoblastic and often blastoporphyritic and poikilitic. Besides
this the reaction hornblende has given rise to a fabric similar to the primary
ophitic (Plate vi, figs. 4, 5). The grainsize varies in the different specimens
from 3 mm. to less than 1 mm. The minerals composing these rocks are plagio-
clase, augite, hypersthene, brown hornblende, magnetite, ilmenite, chlorite, epidote,
uralite, apatite, (?)rutile, and a zeolite.

In the blastoporphyritic types the plagioclase phenocrysts are tabular, whilst
that of the groundmass forms small laths or is partly granular. Many of the
felspars show an inverted zoning, with a later outer rim of more acid plagioclase.
The acid core and selvage are of basic andesine and the middle zone consists of
basic labradorite (Ab,.An,,). The addition of more acid felspar from the invading
magma is also indicated by a peculiar mottled appearance of some of the larger
tabular crystals. As in the pyroxene-gabbros, the plagioclase shows a charac-
teristic clearing and the development of minute inclusions, and occasionally
schiller inclusions may be seen within the tiny inclusions of pyroxene in the
felspar. Besides this clear felspar there is a little cloudy plagioclase sometimes
developed. Under the low power objective this appears as a greyish cloudiness
which may be resolved under the high power into minute dark brown dots similar
to, though much smaller than, the tiny inclusions of iron ore in the cleared felspars.
Clouding and granules are often present together in the same felspar crystal
and the clouding may be confined to definite zones or lamellae of the plagioclase.
The most noteworthy feature of the felspar of these rocks is its apparent reaction
with the invading magma to form brown hornblende (Plate vi, fig. 4). This
process was noticed as an incipient change in some of the specimens of the last
group, but among the hornblende-pyroxene-gabbros the process is far advanced
and large sheets of basaltic hornblende appear to be threaded in between the
plagioclase crystals and often enclose them. Where hornblende has become an
important constituent of the rock, it may be seen encroaching along cleavage
planes and forming embayments in the felspar.

Augite and hypersthene are developed in subidioblastic columns or small
xenoblasts. In a few cases the larger columnar pyroxenes are grouped, indicating
a glomeroporphyritic fabric in the original rock. In one slide these groups show
a rosette-like arrangement as observed by Dr. C. E. Tilley (1921) in some of the
metadolerites from Hyre’s Peninsula, South Australia. Schiller inclusions or a

132 PETROLOGY OF THE HARTLEY DISTRICT, li,

heavy discharge of secondary magnetite, often in a dendritic pattern, sometimes
mask the nature of the original pyroxene. Both augite and hypersthene are
flecked with brown hornblende and are usually surrounded by a wide reaction-
rim of the same amphibole. The rounded and corroded appearance of the
pyroxene crystals testifies to the origin of the brown hornblende. In a few slides
masses of iddingsite are pseudomorphing the hypersthene. This change is probably
deuteric and has been recorded as such in the hypersthene-andesites of Blair
Duguid by W. R. Browne and H. P. White (1926). The basaltic hornblende
shows undoubted evidence of a secondary origin, and apparently arises as a
result of reaction between the invading magma and both plagioclase and pyroxenes.
It forms large irregular crystals up to 5 mm. across, and is highly poikilitic,
enclosing partly dissolved and corroded crystals of the minerals from which it
has originated. The absorption is Z> or = Y>X. In a few rocks situated close
to the quartz-mica-diorites, the hornblende is a less basic type and is greenish-
brown in colour. In some of the sections an outer border of bluish-green soda-
bearing hornblende has been noted.

The small ellipsoidal bodies, showing a zoned arrangement, and consisting
of granular plagioclase, pyroxene and iron ores, are fairly common amongst
this group of rocks. Occasionally they are surrounded by a reaction-rim of
brown hornblende.

One rather unique specimen of hornblende-pyroxene-gabbro has been collected
in which much of the brown hornblende is idiomorphic. Tiny acicular felspars
are crowded between slightly more basic plagioclase laths and these little needles
are abundant as inclusions in the ferromagnesian minerals.

iv. Hornblendites.

The hornblendites occur in small rounded, lens-like or vein-like masses in
the “reaction’-gabbros, and are particularly abundant in the northern part of
Por. 237, Parish of Lowther, on the property of Mr. Chris Commens, and in the
southern part of Por. 27, immediately north of Por. 237, on the property of
Mr. D. Mitchell.

These rocks have already been described in some detail (Joplin, 1931), and
little further need be said regarding their petrography. It might be pointed out,
however, that although these small ultrabasic masses were previously regarded
as xenoliths, it was suggested, on account of their vein-like appearance, that they
might represent basic segregations. It was further suggested that the rocks are
hybrids on account of the comparatively low specific gravity, relatively acid
felspar (andesine) and the presence of small quantities of quartz. The fact that
a fairly acid mesostasis is present also points to hybridization. The occurrence
of zoned hornblende with a sodic border suggests reaction, and calcite, epidote
and uralite indicate deuteric activity.

v. Diorite-gabbros and hornblende-gabbros.

A group of rocks termed ‘‘diorite-gabbros” was described by the writer (1931),
and it was considered that these formed a reaction-ring about the pyroxene-
gabbros.

The present study on metamorphism, however, has revealed the fact that
these rocks are not so abundant as at first appeared, and many of the rocks
thought to have been diorite-gabbros now prove to have suffered an earlier
recrystallization and can be assigned to the group of hydrothermally altered
gabbros described below.

BY GERMAINE A. JOPLIN. Ue

The rocks that are now considered to be diorite-gabbros occur in isolated
patches among the quartz-mica-diorites and “reaction’’-gabbros, and as their
appearance is often very similar to the contiguous rocks, it is likely that they
are far more abundant than at present appears to be the case.

The name diorite-gabbro has been retained for this group as the rocks have
an ophitic or subophitic fabric and the chief ferromagnesian mineral is green
hornblende such as is common in the diorites. They are, however, decidedly
more basic than the quartz-mica-diorites, and the felspar is a labradorite, which
characterizes the gabbros. It will be shown later that these rocks are possibly
hybrids due to the contamination of the quartz-mica-diorite magma by solid
gabbro, so the name “diorite-gabbro”’ seems quite fitting and satisfactory.

In the handspecimen the diorite-gabbros have the appearance of diorites
and seem to consist of hornblende and plagioclase in about equal proportions.
On Marriott’s Creek and near the head of Hughes’ Creek these rocks have a
rude banding and are slightly richer in felspar.

Under the microscope the diorite-gabbros are seen to have an ophitic and
poikilitic fabric and to consist of plagioclase, brownish-green or green hornblende,
iron ores, occasionally augite, and in a few specimens a little quartz and biotite.
Secondary sphene, epidote, chlorite and kaolin bear testimony to a period of
deuteric activity or hydrothermal metamorphism.

The plagioclase is exceptionally abundant, especially in the banded types.
It is labradorite (Ab,,An,,),* and is occasionally zoned. The centres of the
felspar sometimes show alteration into sericite and kaolin, whilst the outer
borders are quite clear. In one slide the banded nature of the rock is accentuated
by the parallel arrangement of the plagioclase laths.

The hornblende is either green or greenish-brown and forms highly poikilitic
patches about 1-5 mm, across. In the banded types these patches are arranged in
linear fashion. The more idiomorphic hornblende crystals are sometimes
surrounded by a bluish-green sodic border.

In one specimen fresh augite is present as well as hornblende and its poikilitic
and linear arrangement is similar to that of the amphibole.

A little highly poikilitic biotite is present in a few slides, and is usually
associated with (?)lawsonite and chlorite. Quartz is occasionally present as
tiny interstitial grains, apatite is fairly abundant and iron ores, though not
apundant, behave in a similar manner to the ferromagnesian minerals. The
iimenite is sometimes surrounded by grains of secondary sphene.

The hornblende-gabbros also have a sporadic distribution and in the hand-
specimen are dark, heavy, fairly coarse grained rocks consisting of plagioclase and
hornblende.

Under the microscope they have an ophitic fabric and are seen to be
composed of brown or dark brownish-green hornblende, pyroxene (usually
completely uralitized), plagioclase, iron ores and apatite. A little quartz and
biotite has been noted in a few specimens, and hydrothermal activity is evidenced
by the presence of uralitized pyroxene, uralite veins, secondary magnetite, epidote
and (?)lawsonite. In one slide an original hornblende crystal is pseudomorphed
by tiny flakes of biotite and chlorite, which suggests a weak thermal meta-
morphism. Slight pneumatolysis is evidenced by the presence of a little pyrites

*It was previously stated (1931) that the felspar was occasionally as basic as
Ab,,An,,, but these specimens have proved to be hydrothermally altered gabbros.

134 PETROLOGY OF THE HARTLEY DISTRICT, ii,

and tourmaline in a specimen near the boundary fence between portions 27 and
239, Parish of Lowther.

These rocks need not be described in detail, as they are very similar to the
deuterically altered hornblende-gabbros in the centre of the intrusion, and their
close similarity will be commented upon later.

vi. Hydrothermally metamorphosed gabbros.
Both the recrystallized pyroxene-gabbros and the “reaction’’- or hornblende-
pyroxene-gabbros have suffered hydrothermal alteration in sporadically distributed
localities.

(a) Altered pyroxene-gabbros. ‘

Like the recrystallized pyroxene-gabbros, the hydrothermally metamorphosed
types show a great variety of textures, but an earlier partial granoblastic structure
may usually be made out, thus showing that these rocks are altered recrystallized
gabbros.

The mineral constituents are wholly or partly uralitized pyroxene, plagioclase,
magnetite, ilmenite, apatite, chlorite, talcose material, secondary sphene and
sometimes iddingsite, limonite and carbonates.

The original phenocrysts or larger pyroxene crystals are usually heavily
schillerized or crowded with secondary magnetite grains arranged in a dendritic
pattern; but those crystals that have suffered an earlier recrystallization do not
show this discharge of iron ores to the same extent. Frequently the pyroxene is
converted into a pale uralitic amphibole, and uralite veins are not uncommon
(Plate vi, fig. 3). In cases of less extreme alteration the uralite may fringe
the pyroxene. The hypersthene often shows cracks filled with a talcose material,
or may sometimes alter into iddingsite. In a highly altered rock the pyroxenes
show a good deal of carbonation.

The plagioclase evidences an earlier partial recrystallization, sometimes by its
granular appearance, but more often by the presence of the characteristic small
inclusions. A further change is noticeable in most of these rocks, namely, a
cloudiness similar to that which has been observed in the “reaction’’-gabbros.
Clouding is often present in the same crystal as the minute inclusions and seems
to be superimposed upon the original clearing. It is usually confined to definite
zones of the felspar—most frequently to the sodic zones. It may also be confined
to certain twin lamellae. Under the high power objective the clouding is resolved
into tiny brown dots, smaller than those associated with the ferromagnesian
granules of the partially recrystallized felspars. The dots suggest iron ores—
possibly ilmenite. The plagioclase may also show streaks of chlorite and in a
type that has suffered extreme alteration it is sericitized and kaolinized.

Ilmenite is usually surrounded by granules of secondary sphene, and apatite
is a little more abundant in these rocks.

(b) Altered hornblende-pyroxene-gabbros.

There is little need for a detailed description of the altered hornblende-
pyroxene-gabbros since they are essentially similar to the “reaction’’-gabbros, and
the hydrothermal changes are similar to those that have just been described for the
altered pyroxene-gabbros. One further change, however, is worthy of note.

As in the “reaction’-gabbros, the brown hornblende often borders the
pyroxenes, and in several of the hydrothermally altered types this reaction-rim
is entirely altered into a mass of pale green actinolite fibres (strahlstein). The

5

BY GERMAINE A. JOPLIN. 13
enclosed cores of pyroxene are heavily schillerized, and the talcose form of
alteration seems more common among the hypersthenes in this group of rocks.
Augite shows alteration into uralite, chlorite and carbonates.

vii. Quartz-mica-diorites.

A fairly detailed description and an analysis of these rocks have already
been published, so little further need be said regarding them.

It has been pointed out that the quartz-mica-diorites form a ring-like outcrop
about the gabbros and, though a little quartz and biotite are present, they are
fairly typical diorites in the handspecimen.

Under the microscope the rocks are somewhat porphyritic in plagioclase,
which is usually zoned. In a single crystal these zones have been observed to
range from Ab,,An,, to Ab,sAny. Several specimens show small inclusions within
the felspar and at first sight they appear similar to those described as charac-
teristic of the partly recrystallized felspars. Examination under high power,
however, reveals the fact that these inclusions are mainly biotite and magnetite,
the former often arranged with their longer axes parallel to the elongation of
their host (Joplin, 1931), and it would thus seem that their origin was different
from that of the granules of the recrystallized felspar. Nevertheless, the
possibility of a similar origin cannot be entirely disregarded and it is possible
that some of the “phenocrysts” are xenocrysts caught up by the quartz-mica-
diorite magma after the shattering and stripping of the outer margin of the
gabbro. The large blocks of pyroxene-gabbro, etc., within the diorite mass
testify that such a tearing away of the selvage of the gabbro did take place.

The deuteric alteration of the quartz-mica-diorites needs a little further
emphasis, and it should be pointed out that the late-emagmatic solutions were
fairly rich in potash. The chief deuteric minerals are sericite, saussurite, kaolin,
muscovite, carbonates, a-zoisite, epidote, chlorite, (?)lawsonite, and secondary
sphene.

In one specimen from Marriott’s Creek a large crystal of plagioclase about
4 mm. in diameter has a clear outer rim showing two cleavages, and the interior
is occupied by a mass of saussurite associated with muscovite. Blades of
muscovite are arranged approximately at right angles to one another and parallel
to the cleavages in the outer rim of felspar. These appear to be threaded through
the granular saussurite which consists of abundant a-zoisite, carbonates and a
little albite. An inclusion of hornblende shows epidotization and a change into
fibres of pale secondary hornblende. There is no reason for assuming this white
mica to be the sodic variety—paragonite—and it has been demonstrated else-
where (Browne and White, 1926, 1928) that potash solutions ane active deuteric
agents,

vill. Hedenbergite-bearing gabbros.

Only two specimens of hedenbergite-bearing gabbro have been collected, one
on Hughes’ Creek and the other on Marriott’s Creek. Their relations to the
surrounding rocks have not been satisfactorily ascertained, but in both cases
they are close to the contacts. On Hughes’ Creek the rock is adjacent to the
southern margin of the intrusion, and on Marriott’s Creek the presence of a mass
of quartzite at approximately the same level and 16 chains to the north-west
indicates the proximity of the roof.

In the handspecimen the two rocks are quite dissimilar. The Hughes’ Creek
type is a very coarse sage-green rock consisting of plagioclase and hedenbergite.

136 PETROLOGY OF THE HARTLEY DISTRICT, il,

The Marriott’s Creek variety is a fine grained, dark olive-green hornfels with a
specific gravity of 3-008. Little can be made of its composition in the hand-
specimen, but small aplitic veins and groups of tiny dark green resinous acicular
crystals may be discerned.

Under the microscope the Marriott’s Creek type can be seen to consist of
three regions and shows recrystallization: (1) A region very rich in pyroxene
and its alteration products; (2) a pyroxene-granulite with a little quartz; and
(3) comparatively coarse quartz-felspar veins.

(1).—These areas consist of abundant crystals of green non-pleochroic
pyroxene which is poikilitic towards tiny grains of quartz and clear untwinned
plagioclase. These crystals usually measure about 0:5 mm. and are flecked
with, surrounded by, or entirely replaced by, chloritic material, and along veins
by a secondary amphibole. Tiny dark, indeterminate inclusions are numerous,
and in these areas small subidiomorphic crystals of sphene are abundant.

(2) —These areas are similar to those described above, but the pyroxene
crystals measure only about 0:07 mm. The plagioclase is zoned and very basic,
the outer rim of the crystals being a little less so. The inner core is sometimes

' sericitized.

_ (3).—The felspar forms idiomorphic laths 0-75 mm. in length and is andesine.
Carlsbad twinning is common and albite and pericline twinning sometimes present.
A little orthoclase is associated and allotriomorphic quartz grains are fairly
abundant. A few pyroxene crystals appear to be torn off from the adjacent
granulite.

The Hughes’ Creek type also consists of three regions: (1) a coarse area
consisting essentially of hedenbergite and basic plagioclase; (2) a region composed
of small grains of the same minerals; and (3) vein-like areas of small, acicular,
more acid felspars associated with granular knots of pyroxene and fairly large
grains of quartz. Quartz veins are also present. As in the hornfels type
(Marriott’s Creek) sphene is abundant and iron ores typically absent.

(1).—The pyroxene is pale greyish-green and forms large subidiomorphic
columns up to 6 mm. in length. Both simple and multiple twinning are present,
and the extinction measures 414°. Diallage cleavage and occasionally a sahlite
striation are present and the centres of many of the crystals are clouded with a
minute dust, possibly the cut ends of schiller inclusions arranged along the
sahlite direction. Slight flecking by brown hornblende and chlorite is common
in the centre of the crystals and one is entirely pseudomorphed by a dendritic
arrangement of iron ores (apparently ilmenite), in a mass of chlorite and tiny
granules of sphene, felspar and quartz. The plagioclase forms large subidio-
morphic to allotriomorphic grains. A few grains show slight evidence of
clearing upon which is superimposed an alteration into kaolin, sericite and
carbonates. Sphene forms interstitial grains up to 0-6 mm.

(2) —This region has a grainsize of about 0-1 mm. and the fabric is
granular. Plagioclase, green pyroxene and sphene are the chief minerals. These
areas appear to have suffered recrystallization.

(8).—These are vein-like areas and are made up of more acid acicular or
lath-shaped plagioclase crystals, which sometimes show zoning, in association
with the above mentioned minerals.

137

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138 PETROLOGY OF THE HARTLEY DISTRICT, ii,

PETROGENESIS.
i. Possible Origin of the Various Rock Types.

Many petrologists, who have studied contact metamorphism, J. Barrell (1907),
V. M. Goldschmidt (1911), E. P. Dolan (1923), H. H. Thomas and E. B. Bailey
(1924), and others, have shown that two stages of metamorphism may often be
recognized: first, a thermal stage or recrystallization produced by heat alone;
and second, a phneumatolytic, metasomatic or hydrothermal stage produced by the
percolation of hot fluids.

The foregoing petrographical descriptions have served. to show that both
these types may be recognized at Hartley, together with a further change, namely,
reaction or hybridization. Table I shows the chief petrographical evidence for
the recognition of these processes among the main rock types.

Reference to this table shows that the chief petrographical reasons for the
assumption that the hornblende-pyroxene-gabbros are hybrids are based on
phenomena which might be attributed to reaction in the normal course of crystal-
lization. There is evidence, however, that the rocks had solidified and become
partly recrystallized before reaction took place. The reacting solutions, there-
fore, must have come from the quartz-mica-diorite magma. The hybrid origin
is further supported by the field evidence, since the “reaction’’-gabbros surround
a partly recrystallized core of gabbro devoid of hornblende, and are themselves
surrounded by the quartz-mica-diorites which are rich in hornblende. Chemical
evidence also lends support to this conclusion.

The deuterically altered hornblende-gabbros, which occupy a small area in
the centre of the Cox’s River Intrusion, have suffered no thermal effects. They
may represent, therefore, either a remnant of the original gabbro intrusion, that
has been in no way affected by the quartz-mica-diorite, or the central portion
of the original intrusion, which was still unconsolidated at the time of the later
intrusion. It is evident, however, that reaction took place in the normal course
of crystallization, and that this was followed by a deuteric period as an end-phase
in the consolidation of the gabbro. The hornblende-gabbros were possibly a little
less basic than the rocks that surrounded them; that is, as on Moyne Farm, the
more acid rocks occupied the centre of the intrusion. The occurrence of slightly
more acid plagioclase, and sometimes a little quartz and biotite in the hornblende-
gabbros points to some differentiation having taken place in the original gabbro
magma.

After this differentiation of the gabbro and after the rock had completely
consolidated, with the possible exception of the central core, the mass was encircled
by a quartz-mica-diorite magma, and partly metamorphosed.

It is evident that the first stage of metamorphism, that is, the thermal
stage, was responsible for the partial recrystallization of the whole of the gabbro
mass except the small central core, which either remained unaffected or was as
yet unconsolidated. If the former be the case, then the thermal effect extended
a distance of 10 to 30 chains from the contact.

Immediately following upon the recrystallization, and before the hydrothermal
stage of metamorphism set in, a certain amount of reaction took place between
the solid gabbro and the fluid quartz-mica-diorite, thus giving rise to the
hornblende-pyroxene-gabbros or “reaction’’-gabbros. It is significant that these
rocks have been partly recrystallized, and always form an outer, though very
irregular, border about the pyroxene-gabbros. Most of the rocks of this group
appear to be of the nature of acidified gabbros, but one specimen (p. 132) might

BY GERMAINE A. JOPLIN. 139

be termed more correctly a basified diorite. This rock contains idiomorphic
brown hornblende and two generations of plagioclase, the later being the more
acid and of an acicular habit. The larger felspar crystals and. pyroxenes
surrounded by reaction-rims of brown hornblende possibly represent the original
constituents of a gabbro, with a grainsize of about 1 mm., which has been
completely disrupted. The original minerals have been thus scattered and some
solution of the basic material has taken place giving rise to a sufficiently basic
liquid to produce brown hornblende on crystallization. The strewing of the
original minerals has allowed space for the brown hornblende to take on an
idiomorphiec form. The acicular felspar possibly crystallized from the substance
of the more acid magma, although its presence in the pyroxene places a grave
objection in the way of this interpretation.

Reaction and the production of brown hornblende from pyroxene and plagio-
clase, together with the probable deposition of hornblende from the substance of
the invading magma, apparently took place more readily along certain lines of
weakness in the hornblende-pyroxene-gabbro, possibly owing to higher tempera-
ture conditions produced by the concentration and escape of volatile constituents.
Vein-like segregations of hornblende and a mesostasis of slightly more acid
material from the fluid magma were thus formed, giving rise to the horn-
blendites.

At certain localized points, possibly where there was an escape of volatile
constituents, superheating occurred and the solid gabbro entirely or partly
dissolved in the more acid magma, giving rise to diorite-gabbros and hornblende-
gabbros. The former are basified diorites and the banded types point to a °
drawing out of the incompletely dissolved gabbro. The hornblende-gabbros repre-
sent acidified gabbros and bear a very close resemblance to those in the centre
of the complex. In fact, it might be argued that the central mass represents a
larger outcrop of these sporadically-distributed hornblende-gabbros, and just
chances to occupy a central position. The central gabbros, however, show
incipient recrystallization and are enveloped by the pyroxene-gabbros, which
are free from the sporadically-distributed hornblende-gabbros. It would thus
appear that the central mass antedated the quartz-mica-diorite invasion. The
two hornblende-gabbros might be expected to have a similar composition, as the
central gabbro possibly arose as the more acid differentiate of the original gabbro
magma, whilst the scattered patches of hornblende-gabbro originated as products of
the reaction between the more basic differentiates of this early magma, and a later
more acid magma.

Whilst these processes were in progress, the quartz-mica-diorites were crystal-
lizing. Near the outer margin the temperature was falling rather rapidly, and
the composition of the magma constantly changing, as the highly-zoned felspars
show. It is probable that the first consolidated layers formed a non-conducting
envelope about the intrusion, and it is for this reason that the various processes
of reaction and hydrothermal alteration have advanced so far.

Reference to Plate v shows that isolated patches of recrystallized pyroxene-
gabbro ovcur within the quartz-mica-diorite and fairly close to the margin of the
intrusion. These evidently represent fragments that were rifted off from the
gabbro mass and made their way towards the margin whilst the quartz-mica-diorite
magma was still fluid. They were thus recrystallized, but owing to their
proximity to the contact they were caught up in the rapidly cooling and solidifying
diorite and the temperature was not maintained for a sufficient length of time

140 PETROLOGY OF THE HARTLEY DISTRICT, ii,

for reaction to take place. The occasional isolated patches of hornblende-
pyroxene-gabbro evidently originated in a similar manner, but in this case
temperature conditions favoured some reaction.

As an end-phase in the consolidation of the quartz-mica-diorite magma a
period of deuteric activity set in. This is evidenced in the diorites by the presence
of epidote, sericite, saussurite, (?)lawsonite, etc. Accompanying this autometa-
morphism of the quartz-mica-diorites, aqueous, aqueo-siliceous and other deuteric
solutions made their way along lines of weakness in the gabbros, and the rocks
were thereky hydrothermally altered. It might be noticed that the chief product
of this stage of metamorphism is uralite, and it seems likely that the aqueous and
aqueo-siliceous solutions that give rise to this mineral were more abundant, more
active and more mobile than the other deuteric solutions, and were thus able to
penetrate the neighbouring rocks.

Gabbro Magma.

@ More Basic Differentiate (4)More Acid Differentiate
{ Hornblende-Gabbro)

Quartz-mica-diorite Magma

Recrystallized
Pyroxene-Gabbro

“ Reaction’~Gabbro or
Hornblende-Pyroxene-Gabbro

H ydrothermally Altered
“ Reaction’~ Gabbro

Deuterically Altered

Quartz-mica-diorite

Hydrothermally Altered
‘Pyroxene-Gabbro

TABLE II.

Table II represents a diagrammatic attempt to show that the chief rock types
have been produced as a result of differentiation followed by the three types of
metamorphism due to the quartz-mica-diorite intrusion: (1) Thermal meta-
morphism; (2) Reaction; (3) Hydrothermal metamorphism.

The hedenbergite-bearing gabbros appear to have quite a different origin
from the types discussed above, and seem to have resulted from the addition of
lime and silica to the gabbro magma. Such a combination could occur only in a
sediment, and the presence of abundant calc-silicate hornfelses in the aureole
points to contamination by sediments.

BY GERMAINE A. JOPLIN. 141

ii. Physico-chemical Discussion.

The assumption that some of the Hartley rocks are hybrids calls for some
justification on physico-chemical grounds.

It has been shown that field occurrence and petrography point to four groups
of rocks of mixed origin:

(1) The hornblende-pyroxene-gabbros, which are apparently due to reaction
between solid gabbros and the fluid quartz-mica-diorite magma, and therefore
represent acidified gabbros. One specimen, described on pages 132 and 138, might
be regarded as a basified diorite.

(2) The hornblendites, which are segregations of “reaction” brown horn-
blende that has erystallized in a mesostasis of more acid material.

(3) The isolated patches of hornblende-gabbros and diorite-gabbros, which
appear to be true hybrids, and represent complete solution of the solid basic rock
in the more acid magma.

(4) The hedenbergite-bearing gabbros that are apparently due to the assimila-
tion of sediments.

It is obvious that different physical and chemical conditions were responsible
for the formation of these different types, whose divers origins have already
been discussed. It has been suggested that (1) and (2) are due to reaction, and
(3) to solution, whilst (4) may be accounted for by the assimilation of sediments.
It is now proposed to discuss the physico-chemical conditions responsible for (a)
reaction, (b) solution, and (c) contamination by sediments.

‘ (a) Reaction or partial hybridization.

Dr. N. L. Bowen (19220) has shown “that the solution of a silicate in a
magma is usually accompanied by a large absorption of heat, probably of the
order of magnitude of the heat of melting”.

Bowen has also shown that if the melting point of the solid phase be lower
than that of the fluid in which it is immersed, complete solution will take place.
As an example of this, he takes the case of a granite inclusion in a basaltic
magma. The granite is supersaturated with all higher (i.e., more basic) members
of the reaction-series (Bowen, 1922a) and complete solution will take place in
the basaltic magma. If, however, a basic inclusion be immersed in an acid
magma, the fluid can not dissolve it, but may react with it and convert it into
such members of the reaction-series with which the fluid is saturated. It is
evident, therefore, that without the aid of superheat a solid gabbro cannot
go into solution in a diorite magma.

The basic felspars and the pyroxene of the recrystallized-gabbros at Hartley
could not dissolve in the quartz-mica-diorite magma, but they were able to react
with it and give rise to zoned and mottled felspar and reaction-rims of brown
hornblende, since green hornblende and andesine are the members of the reaction-
series with which the diorite magma was saturated.

Moreover, Bowen considers that the magma may not only react with solid
gabbro, but may deposit, from its own substance, the minerals with which it is
saturated. In the case of the hornblendites, which possibly represent segrega-
tions of the “reaction” hornblende along cracks in the gabbro, the borders of
greenish hornblende and the acid mesostasis were probably deposited from the
substance of the invading magma.

Bowen (1915, 1922a) has shown that reaction takes place in a magma during
the normal course of crystallization, and as it will be shown below that the

B

142 PETROLOGY OF THE HARTLEY DISTRICT, ii,

parent magmas at Hartley are consanguineous, these so-called partial hybrids
might be regarded as normal differentiates of an original parent magma. The
gabbros, however, were undoubtedly solid before the intrusion of the later more
acid partial magma, and the progress of differentiation was thereby interrupted
and physical conditions changed. H. H. Thomas and BH. B. Bailey (1924) point
out that an internal migrating partial magma, still in the fluid state, will react
with an earlier separated solid phase, and they distinguish between this process
and that of true hybridization when an independent magma acts upon an already
consolidated rock. These authors also say, ‘It will be seen that the two processes
have much in common, and in extreme cases the results may be indistinguishable”.
The Hartley rocks seem to be a case in point, but the evident temperature change
rather favours their being called hybrids.

(0) Solution.

A complete solution has been postulated for the origin of the isolated out-
crops of the hornblende-gabbro and diorite-gabbro, since the fabric of both is
ophitic, the composition is intermediate between gabbros and quartz-mica-diorites
and there is no evidence of recrystallization. Dr. Bowen admits the possibility
of a complete solution of a basic rock in an acid magma where there is super-
heating, but points out that superheating is not a common occurrence.

It has already been indicated that the Hartley gabbros and diorites are
comagmatic, and it is fairly evident that the gabbros were still hot and the
surrounding country rocks considerably warmed up at the time of the quartz-
mica-diorite intrusion. It is even conceivable that the diorites may have been
injected at a temperature above that of the fusion point of the gabbros at the
reduced pressure of their higher position in the earth’s crust. The partial
recrystallization of the gabbros, however, shows that this was not the case, but
we can at least admit a temperature not far below the fusion point, and little
superheat would be required to attain it. Moreover, when this temperature was
acquired the heated country rocks would prevent a rapid dissipation. The extra
heat required to raise the gabbros to the fusion point might be supplied by
escaping gases, and the localized occurrence of the hornblende-gabbros and diorite-
gabbros favours this explanation. Solution would possibly be aided by the high
iron content of these rocks acting as a flux, and once solution had commenced no
further superheat would be required, since, as indicated by Daly (1914), the
experimental work of Petrasch and Doelter has shown that the melting point
of a mixture of rocks of different composition is lower than that of either of the
separate rocks.

The fact that both hornblende-gabbros and diorite-gabbros arose in this way
points to different temperature conditions in different parts of the mass. The
more basic hornblende-gabbros indicate that a greater proportion of gabbro went
into solution, and it would thus seem evident that the temperature at which
these rocks consolidated was greater than that at which the diorite-gabbros
solidified.

The banded diorite-gabbros possibly represent the incomplete solution and
softening and drawing out of the gabbros (Harker, 1904, p. 117) or else indicate
ail immiscibility of the liquid magmas.

(c) Contamination by Sediments.
Bowen and his co-workers at the Carnegie Institution (Bowen, 1922b) have
shown experimentally that the addition of lime to a basaltic magma is exothermic,

BY GERMAINE A. JOPLIN. 143

and no very great modification of the magma is possible without superheating.
They show, however, that reaction may take place and that a more calcic plagio-
clase and pyroxene may result. This appears to be what has happened in the
ease of the hedenbergite-bearing gabbros at Hartley, and the quartz veins and
interstitial quartz grains possibly represent assimilated quartz, which has readily
dissolved in the basic magma and separated out as an end phase. Sphene has
arisen in place of ilmenite as a result of the assimilation of lime.

The recent work of Dr. C. E. Tilley in Ireland (1929, 1931) has shown that
very important modifications of the magma actually do occur, but he agrees with
Dr. Bowen in regarding the effects as limited, and this is again borne out by the
very localized outcrops of these contaminated rocks in the Cox’s River Intrusion.

iii. Origin of the Ellipsoidal Bodies.

Four possible explanations of the origin of these bodies suggest themselves:
(1) An original amygdaloidal or miarolitic structure. (2) An original orbicular
structure. (3) Contamination followed by recrystallization. (4) Recrystalliza-
tion of original zoned hornblende crystals.

Dr. Harker (1904, p. 51) has shown that comparable structures among the
recrystallized basalts of Skye represent originally zoned infillings of amygdules,
but the sporadic distribution and the nature of the rock tends to discount such
an origin in the case of the Hartley ellipsoids.

In considering the second possibility it is pertinent to summarize the chief
characteristics of an orbicular structure (Lawson, 1904; Iddings, 1909; Cole, 1916).

1. The orbicular facies of any rock mass is confined to a definite area which
grades out into the normal rock. The orbicules are not sporadically scattered
through the normal facies of the rock.

2. The distances between the centres of these bodies are fairly uniform.

3. The orbicules are usually, though not always, spheroidal.

4. They are usually large, the size being often 9 cm. to 10 cm., or even
larger.

5. There is nearly always a distinct radial arrangement of the constituent
minerals,

6. The orbicules show concentric zones consisting either exclusively of a
single mineral or of a certain definite mineral assemblage.

7. The centre of the spheroid is usually slightly coarser in grainsize than
the zone surrounding it.

8. The centre usually consists of, or is very rich in, felspar.

With regard to the Hartley gabbros the distribution of the rocks containing
these bodies is somewhat sporadic, although their occurrence always seems to
be confined to the north-western part of the mass (see Plate v). Moreover, the
number of bodies in a single slide is very variable, showing that they are
unequally spaced. Again it is evident that the bodies in the Hartley gabbros
are ellipsoids, never spheroids; and their size never exceeds more than 10 mm.,
that is, they are about one-tenth the size of a normal orbicule. A _ parallel
arrangement has been noted in a few sections, but never a radial, and more-
over, the parallelism is not always present. The last three conditions tabulated
are certainly fulfilled by the Hartley ellipsoids, but the bulk of evidence tends
to show that they do not represent an original orbicular structure.

Orbicular rocks exhibiting some rather remarkable features have recently
been described from Alderney by S. R. Nockolds (1931) and A. E. Mourant (1932).

144 PETROLOGY OF THE HARTLEY DISTRICT, ii,

The following is a summary of their work and conclusions:

1. The orbicules may be widely scattered.

2. Though apparently fairly uniform for any particular locality the size
of the orbicules may vary from about 20-40 mm. (Nockolds) to about a foot
(30 cm.) in length (Mourant).

3. The orbicules are usually flattened spheroids.

4. They are associated with basic xenoliths of a comparable size.

5. The radial structure is absent or poorly developed.

6. The orbicules are slightly different in mineral composition from the normal
rock enclosing them, and bear evidence of having suffered hydrothermal action.

7. The orbicules are surrounded by material which is coarser in grain than
the normal rock and resembles the orbicule more closely in mineral composition.

8. Both authors agree that the orbicular structure is due to contamination.

The small size of the orbicules examined by Nockolds, the shape, and the
frequent absence of radial structure is in accord with the Hartley type, but at
Hartley there is evidence neither of a mineralogical difference between the
ellipsoids and the enclosing rock, nor of the presence of assimilated xenoliths.
The Hartley ellipsoids might be accounted for, however, by postulating
sporadically-distributed xenoliths of number insufficient to produce an orbicular
structure. If these were completely assimilated by the gabbro, it is possible that
minor mineralogical and textural differences might be obliterated by recrystalliza-
tion. The fact that the ellipsoids are completely granoblastic whilst the remainder
of the rock shows relict structures, however, presents a difficulty in this inter-
pretation.

The possibility of the ellipsoids being original zoned hornblende crystals may
now be considered.

1. The sporadic distribution is not altogether out of harmony with this
view.

2. The ellipsoids occur only in the recrystallized types, although this is no
very definite criterion, since they are never abundant and only a few specimens
of the unaltered gabbros are available. Furthermore, the unaltered types may
represent a different facies.

3. If a section of an ellipsoid be viewed macroscopically against a light back-
ground, the shape suggests that of a section of hornblende.

4. It is believed that hornblende was an original constituent of the gabbros,
and is now represented by granules of augite and hypersthene, and the basic
outer zones of the plagioclase. It is possible, therefore, that large zoned horn-
blende crystals may have broken down in similar fashion and that the various
mineralogical zones of the ellipsoid correspond to composition zones in the
original hornblende crystal.

5. There is some chemical evidence for this assumption (p. 152), although it
is difficuit to account for the textural differences of the various zones.

6. The presence of comparatively large stable augite crystals suggests that
the original rock may have been porphyritic in ferromagnesian minerals, and
that the recrystallization of augite alone was not responsible for the present
mineral assemblage in the recrystallized gabbros. Moreover, the size of the
smaller ellipsoids is quite comparable with that of the augite crystals, although
no actual hornblende phenocrysts have been noted in the few unaltered hornblende-
gabbros examined. Text-figure 2 shows a large augite crystal surrounded by a

BY GERMAINE A. JOPLIN. 145

reaction-rim of hornblende and on recrystallization this would probably produce
zoned mineral assemblages.

7. The parallel arrangement, which is not present in every section, may
represent a particular direction in the original crystal.

8. The cross-like arrangement of the coarser central zone in one of the
ellipsoids may represent a kind of original hour-glass structure in the hornblende.

In many respects this last explanation of the origin of the ellipsoids seems to
be the most likely, but at the same time the possibility of contamination followed
by recrystallization cannot be entirely dismissed.

iv. Consanguinity of the Parent Magmas.

The consanguinity of the gabbros and quartz-mica-diorites is indicated by
three criteria, namely, field relations, mineralogical constitution and chemical
composition.

(1) Field Relations—In the opinion of Dr. Harker (1904, p. 169) the very
intimate association of two rocks in the field is indicative of their close genetic
relation. The Hartley rocks are closely associated both in time and space. The
fact that the gabbros were still at a high temperature, and the central core
possibly unsolidified when the diorites were injected points to their proximity in
“time’; and that the second intrusion encompassed the first shows their relation
in “space”.

(2) Mineralogical Constitution—The foregoing descriptions of the rocks in
this paper, together with those published in 1931, have served to indicate that
there is a close mineralogical relation. The unaltered core of hornblende-gabbro
at the centre of the Cox’s River Intrusion compares closely with the intermediate
types that have suffered no metamorphism. In all these types hornblende is the
dominant femic mineral, and in the recrystallized gabbro it seems likely that
the greater part of the rhombic pyroxene has been produced as a result of the
thermal metamorphism of brown hornblende. Biotite is also very characteristic
of this rock series and is found in small quantity in the unaltered hornblende-
gabbros.

(3) Chemical Composition—kReference to Table III will show that the
composition of the recrystallized pyroxene-gabbro (column iii) is very close to,
though slightly more basic than, that of the hornblende-pyroxene-gabbro (column
iv), which was erroneously believed to represent the composition of the original
gabbro magma (Joplin, 1931). It has already been indicated that the pyroxene-
gabbros have eucritic affinities, and this finds chemical support in the analysis
of the recrystallized pyroxene-gabbro in column ii, which represents a slightly
more leucocratic phase.

Although the writer has already shown (1931) the relation of the horn-
blendite, hornblende-pyroxene-gabbro, diorite-gabbro and the Cox’s River type of
quartz-mica-diorite to the intermediate and acid members of the series, it is
deemed necessary to repeat the analyses of these rocks (Table III), and to place
the recently analysed recrystallized pyroxene-gabbros on a variation-diagram with
those types that have suffered no apparent hybridization (Text-fig. 3). This
diagram, therefore, represents the serial differentiation of the magma in an
intercrustal reservoir and a simple secondary differentiation of the partial magmas.
It is evident that this variation-diagram has affinities with the linear type. Some
smoothing of the curves has certainly been necessary at the basic end, but from
this it is apparent that there has been secondary differentiation among the gabbros

ll,

PETROLOGY OF THE HARTLEY DISTRICT,

146

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BY GERMAINE A. JOPLIN. 147

themselves. If reference be made to a variation-diagram of any typical calcic
province plotted in such a way that the silica percentages are used as abscissae
and the other oxides as ordinates, it will be seen that the oxides, with the
exception of the alkalis, usually fall upon straight lines when the silica percen-
tage is above 52. According to Holmes (1921) the Hartley series belongs to a
calcic complex, since the lime curve bisects the vertical between the alkali curves
at a silica percentage of 663, although it will be shown later that this series

3 2
5 5
3 = ‘ z 2
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a3 oO ones 5
ce E a4 Beacon é
S x 7 as 2 = uo re
no N OH 5 ) Pam o
Ser » nS =) o) ar was) oO
SS a 5 a (=! Q ° — ras)
oot fo] _ ~ —
o * 3 oo 5 © 5 2 ° =
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t Sep Se
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semereveer Nn ge
=
Seo), 45 50 55 60 65 70 15
—__-
Text-fig. 3.

exhibits certain peculiarities. The fact that the curves are approximately straight
between 54:37% and 76:94% of silica is to be expected, but that they continue to
remain fairly straight as far as a silica percentage of 44-52 calls for comment.
Harker (1909), in giving a mineralogical interpretation of a sub-alkaline variation-
diagram, has pointed out that anorthite contains 37% of alumina and the other
felspars less. As the Hartley gabbros, particularly the leucocratic types, have
eucritic affinities the alumina percentage is considerably higher than that of
normal gabbros with a similar silica percentage. The alumina, therefore, follows
a gentle curve towards the quartz-mica-diorite instead of abruptly rising.
Similarly since there is an antipathetic relation between AI.0, and Fe,0,, the iron
oxides, instead of abruptly descending, come down on a very gentle curve. It is
obvious that, had the leucocratic phase of the gabbro, with its high alumina and

148 PETROLOGY OF THE HARTLEY DISTRICT, 1i,

lew iron oxides, been considered alone, the alumina and iron oxide curves could
have been drawn as straight lines.

A second variation-diagram (Text-fig. 4) has been plotted to show the rela-
tions of the hybrid rocks to the parent magmas. The tonalite is included in this
diagram since it is a differentiate of the Moyne quartz-mica-diorite, whose
composition is possibly close to that of the parent quartz-mica-diorite magma. It
will be seen that with very little smoothing the curves are typical of the basic

Quartz-mica-diorite

Pyroxene-gabbros
Quartz-mica-diorite

Hornblendite
Recrystallized
Diorite-gabbro

= “Reaction’’-Gabbro
Tonalite

>}

ae 45 50 55 é0

Text-fig. 4.

end of a calcic or sub-alkaline series. The fact that the secondary differentiation
of the gabbros is readily smoothed out and that the hybrids take their place upon
the normal curves further supports the genetic relation of the two reacting types
and emphasizes the slight distinction between normal differentiates and hybrid
rocks arising from cognate parents, when free diffusion has been able to take
place.

It seems evident that at least two processes operated in the differentiation of
the Hartley series—gravity separation and assimilation. The writer has previously
indicated (1931) that there were possibly three main acts of intrusion from an
intercrustal reservoir—(1) gabbro; (2) quartz-mica-diorite; (3) granite; and
presumably these three types are due to gravity separation. The results recorded
in the present paper indicate that the series between the gabbro and quartz-mica-

BY GERMAINE A. JOPLIN. 149

diorite is the result of hybridization; thus assimilation, though it be concerned
with comagmatic types, is an important factor in giving rise to this rock series.

In an endeavour to show that differentiation does not always take place as a
result of the gravitational separation and the sinking of crystals, C. Fenner
(1926) has reviewed the various theories of differentiation and concludes that a
linear variation-diagram is indicative of the operation of a process other than
gravity-sinking.

With regard to the Hartley series, the types supposed to be due to gravity
separation alone readily take their places upon a somewhat linear type of
variation-diagram, and those that are proved to be hybrids fall upon curves in
direct contrariety to the arguments of Fenner. It is true that the gabbros have
suffered secondary differentiation and, had the normal type alone been plotted,
the linear relation for the basic end of the series would not be so apparent. It
has been pointed out, however, that the linear type of diagram is not uncommon
for a calcic series with a silica percentage above about 52. This being the case,
must the differentiation of all such series be attributed to some cause other
than the gravity separation of crystals? There seems to be a danger in comparing
the diagram of an actual rock series with one of an ideal case such as the
diopside-albite-anorthite series (used by Fenner), which has been derived experi-
mentally under laboratory conditions. The writer does not feel competent to
discuss this question, but at the same time is not prepared to attribute the
differentiation of the whole of the Hartley series to assimilation, or to some
process other than gravity separation.

An examination of Table III and of Text-figures 3 and 4 shows that the
Hartley series have certain chemical peculiarities, and in some cases these can
be interpreted mineralogically. It will be noticed that alumina is exceptionally
high throughout the series, that the potash curve crosses that of soda at a silica
percentage of about 66 instead of in the more normal position of about 68%, and
that in the gabbros potash is extremely low. These facts can possibly be
explained by the eucritic affinities of the gabbros and by the high biotite content
of most of the other members of the series. The alumina would be contained
mainly in the anorthite-molecule in the gabbros, and with the coming in of
potash, in the more acid rocks, would enter into the biotite-molecule. Biotite is
the chief ferromagnesian mineral in the acid rocks of this series and in the
intermediate types it is equal to or even greater than hornblende, whilst it is
present as an accessory in some of the deuterically altered hornblende-gabbros.
Titania is rather low in the gabbros, and Holmes (1921, p. 459) has pointed out
that there is an antipathy between this oxide and normative anorthite.

Reference to other typical calcic series shows that magnesia is usually
higher than ferrous oxide, but at Hartley the reverse is the case. Moreover,
soda is comparatively high for a normal sub-alkaline complex. Washington (1915)
has shown that “soda not uncommonly tends to vary with the iron oxides,
while potash shows similar relations to magnesia’. The first statement seems
to be true for the Hartley rocks, but the relation between potash and magnesia
does not hold.

Besides the high ferrous oxide it will be noticed that ferric oxide is unusually
high and often approximately equal to the ferrous oxide. This can possibly
be explained by deuteric alteration, which is common to most members of the
series.

150 PETROLOGY OF THE HARTLEY DISTRICT, ii,

v. Chemical Changes involved during Hybridization and Metamorphism.

It is not unreasonable to assume that the quartz-mica-diorites on Moyne Farm
and on Cox’s River originated from the same partial magma, and were originally
of the same composition. The Moyne magma differentiated in situ, giving rise
to a central mass of tonalites and an outer fringe of more basic quartz-mica-
diorite. The original partial magma was, therefore, more acid than the Moyne
quartz-mica-diorite, since this latter represents the original type depleted of
some of its more acid constituents by differentiation. Assuming this to be the
case, it is evident that the Cox’s River type has been still more basified, although
no acid differentiate is apparent. The basification has probably been effected by
assimilation of gabbro, and the diorite is thus a hybrid.

According to Nockolds (1931) “reciprocal reaction is the very essence of
contamination’, but unfortunately this is impossible to show in the case of the
Hartley series, since the true composition of the reacting quartz-mica-diorite is
unknown. Reference to Text-figure 4 will show that the hybrid gabbro and
diorite are intermediate in composition between the recrystallized gabbro and
the Moyne diorite and, although the Hartley analyses are not altogether suitable
for such a comparison, the writer should like to point out the danger of a direct
comparison as employed by Foye (1915), Read (1923) and Nockolds (1931).
Merrill (1906), Ransome (1911) and Browne and White (1926, 1928) have already
indicated the inaccuracies arising out of such a comparison, and have shown
that a true comparison can be made only by assuming that some standard
constituent has remained constant, or by comparing unit volumes of the rocks.
By the direct method unit weights are compared and if the two rocks differ in
specific gravity and porosity, then the comparison is made between unequal
volumes of the reacting rocks. Ransome (1911) has discussed this at some
length, and explains that if, for example, one rock be twice the specific gravity
of another, then by the direct method a unit volume of one rock is compared
with half that of the other. It is evident from this extreme case that such a
comparison is invalid.

Unfortunately the comparison of equal volumes can be made only between
two rocks that have remained solid during reaction. Thus the Hartley gabbros,
the parent and the hybrid, may be compared, since the gabbro was a solid and
the “reaction’’-gabbro did not crystallize from solution but was produced as the
result of reaction between the magma and _ solid gabbro. Even if the true
composition of the parent diorite were known, however, it could not be compared
with the hybrid diorite, since the latter crystallized from solution, and it is
impossible to say what volume of the fluid magma reacted with the solid gabbro
to form the hybrid diorite magma.

Table IV shows a comparison between equal volumes of the recrystallized
gabbro and the “reaction’’-gabbro. Both analyses have been re-calculated to 100,
so that equal weights may be the basis of the calculation. It is doubtful whether
this is valid, since it is an assumption that the experimental error is propor-
tionally distributed whilst in reality it may have occurred in the estimation of
a single constituent (Washington, 1919, p. 26). This will serve, however, as a
reasonable basis for an approximate comparison. In the hybrid as compared
with the parent gabbro the specific gravity has been decreased by about 1:6% as
the result of contamination and the analysis of the parent gabbro has therefore
been re-calculated (column E) so that its sum is about 98-4 instead of 100.

BY GERMAINE A. JOPLIN. 151

TABLE IV.
A B C. D | 1D}
|
ele es = = 2 ee ee seas i
SiO, | 44-79 45°31 44°97 45°05 44°21
Al,O; 19-56 19°39 | 19°64 19-28 19-31
Fe,0, 6-01 5-33 6-03 | 5:30 5-93
FeO 7:79 7°81 7°82 7:76 7-69
MgO 6-16 6°93 6:19 6:89 | 6-08
CaO 11-81 11-67 | 11-86 11-60 11-66
Na,O 1-21 1-22 22, ILOPA | | 20
K,0 a4 0 6b 0-06 0°35 0-06 0°34 | 0:06
180) AS es 5 50 0:64 0-69 0:64 0-69 | 0-64
| |
H.O — 0-10 0°08 0-10 0-08 0-10
hiOs ae aie By 1-14 i583 1-14 1-32 1:14
P.O; co 70 30 0-18 0°31 | 0°18 0°31 | 0-18
MnO 0-15 0-17 0°15 0-17 | 0-15
Total ae ee 99-60 100-59 100-00 100-00 98-36
Specific Gravity .. 3-055 3-004. | |
A. Recrystallized Pyroxene-gabbro (Normal Type).
B. “Reaction’’-gabbro.
c. Analysis A re-calculated to 100.
D. Analysis B re-calculated to 100. |
E. Analysis A re-caleulated as explained in text.

Columns D and H, therefore, represent unit volumes of the two rocks, and
if these be compared certain slight gains and losses are apparent. Differences
less than 0:10% have not been taken into account as.it is possible that such are
due to experimental error. It is evident that the gabbro has gained SiO., MgO, K.O,
TiO, and P,0; from the diorite magma, and has lost Fe,0,.

An examination of a series of pyroxene and amphibole analyses (Iddings,
1911; Clarke, 1910) shows that silica, magnesia and potash are higher in the
amphiboles, so it seems possible that these additions from the diorite magma
entered into the composition of the ‘“reaction’’-hornblende.

The chemical changes brought about by metamorphism may now be
considered.

The chief mineralogical changes are the production of rhombic: pyroxene
and more calcic felspar in the recrystallized gabbros, and the formation of
uralite, etc., in the hydrothermally metamorphosed types.

The original mineral composition of the recrystallized gabbros is somewhat
doubtful. The central core of the Cox’s River Intrusion certainly seems to be
a more acid differentiate, and in this rock hornblende is the dominant ferro-
magnesian mineral, so it is likely that this amphibole occurred in the outer
slightly more basic differentiate. The presence of apparently unaltered augite
in some of the recrystallized gabbros, however, indicates that there was at least
some monoclinic pyroxene originally present, and that this was stable under
the thermal conditions. It is also probable that some of the hypersthene is
primary, although the greater part appears to be the product of recrystalliza-
tion. If the primary augite was stable, then brown hornblende is indicated as
the mineral that gave rise to basic plagioclase and secondary hypersthene, augite
and iron ores.

152 PETROLOGY OF THE HARTLEY DISTRICT, ii,

C. E. Tilley (1921), F. L. Stillwell (1923) and W. R. Browne (1927) have
shown that similar changes occur in the breaking down of pyroxenes in metamor-
phosed basic: igneous rocks.

The following equation illustrates the breaking down of the hornblende as
observed at Hartley:

m CaMg,(SiO;); ] (2m-1) CaMg(SiO,), |
2 = \ + 2(m+1)MgSiO, + CaAl,SiO0,
J

MgAl.SiO, MgAl.SiO,
Hornblende Augite Hypersthene Anorthite

In the case of brown hornblende the high iron percentage would be evidenced
by a discharge of iron ores, and such are present in association with the
pyroxenes of the gabbros at Hartley.

The zoned ellipsoids that have been described in the present paper (p. 143)
may thus be accounted for by the breaking down of zoned hornblende consisting
of iron-rich, magnesia-rich and alumina-rich shells.

The chief change produced by hydrothermal metamorphism is the produc-
tion of uralite from pyroxene. This was probably effected by the action of
superheated steam and hot aqueo-siliceous solutions. Deuteric activity among
the quartz-mica-diorites has resulted in the production of lime and potash
minerals. It is possible that muscovite is the result of pneumatolysis connected
with the granites, though it is usually associated with deuteric minerals and may
be paragenetic with them. Only one definite example of pneumatolytic action
has been noted, where a little pyrites and tourmaline occur in a hornblende-
gabbro. The addition of lime (from the gabbros) to the quartz-mica-diorite
magma would possibly account for a late concentration of potash, with the forma-
tion of deuteric lime and potash minerals. The occasional sodic borders of the
hornblende may be accounted for similarly.

vi. Clearing and Clouding of the Felspar.

Apart from the clouding of the felspar due to deuteric products, two types
of alteration have been noted, namely a greyish clouding produced by minute
inclusions, and a clearing accompanied by tiny granular inclusions either
zonally arranged or centrally placed.

A. G. McGregor (1931) has discussed the phenomena of clouding and
recognizes two types—a brown clouding due to the presence of ultramicroscopic
inclusions, and a grey turbidity produced by tiny rod-like or hair-like inclusions
of iron ores. Both types he ascribes to metamorphism, but considers that the
felspar showing the greyish clouding has suffered a slightly higher grade of
thermal metamorphism. Of the tiny granular inclusions he makes no mention,
but states that under the grade of metamorphism which converts amphiboles
into pyroxenes and renders labradorite unstable, the felspar is perfectly clear.

The greyish clouding of the felspar has been observed in several of the
Hartley types—in some of the rocks that have suffered reaction and in some of
the hydrothermally metamorphosed gabbros.

The cleared felspar with the accompanying tiny granules occurs in the
recrystallized gabbros, but the felspar which is quite granular, and has evidently
suffered complete recrystallization, is unaccompanied by granules. In some
instances both greyish clouding and granules are present in the partially recrystal-
lized felspar.

i)

BY GERMAINE A. JOPLIN. 15%

The writer has had the opportunity of examining a number of thin sections
from various localities, exhibiting these phenomena, and it is pertinent to make
brief reference to them.

A collection of metamorphosed dolerites from Broken Hill, kindly lent by
Dr. W. R. Browne, contains perfectly limpid felspar crowded with granules of
pyroxenes and brown hornblende, that are distinctly larger than those observed
in the Hartley rocks. According to Dr. Browne (1927) the granules are some-
times so numerous that the felspar is rendered a grey colour, but it might be
pointed out, however, that this grey colour is quite distinct from the grey cloudi-
ness previously referred to, and that in this case the granules are easily distin-
guished as such. In some of the felspar of these rocks the aggregate of inclusions
is of greater bulk than the thin outer rim of clear felspar. The Broken Hill
dolerites exhibit blastophitic and granoblastic structures, and those felspars that
are completely recrystallized and show no relict lath-habit are perfectly clear
and free from inclusions. The femic minerals in these rocks consist of pyroxenes,
brown hornblende, olivine and iron ores, and the ferromagnesians are heavily
schillerized.

Another interesting collection of microsections was kindly lent by Mr. T. Hodge-
Smith of the Australian Museum. These rocks were collected by Mr. Hodge-Smith
in the Hart’s Range, Central Australia, and appear to represent highly metamor-
phosed dolerites. Several phenomena are represented in the felspar of these rocks.
As in some of the Hartley gabbros the granular, recrystallized felspar is quite
clear and free from inclusions; other rocks partly recrystallized contain granular
femiec minerals indenting the partly granoblastic felspar, and in this case the
felspar shows clearing with tiny granular inclusions or regularly arranged colour-
less or brown plates similar to schiller inclusions. Moreover, the “schiller’ and
granule-bearing felspar is usually accompanied by schillerization of the ferro-
magnesian minerals. Many of these rocks contain felspar that is clouded, and
the grey clouding commonly accompanies the granules or schiller inclusions.
Green hornblende replacing original pyroxenes is the most abundant femic
constituent of these rocks.

Felspars showing colourless or brown schiller plates and, in the opinion of
W. N. Benson (1910), negative crystals, are present in some of the very basic
gabbro xenoliths from the Dundas voleanic neck, and here again the accompanying
pyroxenes are usually heavily schillerized.

H. H. Thomas (1924) has also described gabbro xenoliths from Mull containing
cleared felspar with granular inclusions.

A. Harker (1904) has described metamorphosed basalts from Skye showing
this feature, and the writer has had an opportunity of examining a collection of
these slides in the Geology Department of the University of Sydney.

McGregor (1931) cites a metadolerite from Eyre’s Peninsula, South Australia,
as an example showing cloudy felspar, and the present writer has been able to
examine two microsections from this locality. There appear to be two types
of alteration present—a clearing accompanied by granular inclusions and a
clouding. Dr. C. H. Tilley (1921), in describing these rocks, refers to both of
these phenomena, and McGregor groups them together under ‘clouding’. Accord-
ing to Dr. Tilley, the granules are present only in the relict felspars, whilst those
that are completely recrystallized are quite limpid. These rocks also contain
schillerized ferromagnesians—brown hornblende, augite and hypersthene.

154 PETROLOGY OF THE HARTLEY DISTRICT, il,

A search of the literature dealing with this subject shows that there is much
confusion in the terminology. Some of the Scottish geologists refer to turbidity,
others to schillerization, and McGregor, who has examined many of the Scottish
rocks, is convinced that the phenomena described by these terms are identical
with what he himself terms clouding. The present writer has compared plates
of the Scourie dyke rocks (Teall, 1885, 1888) and examined sections of the
Malchite and Orbite from Melibocus, Odenwald, referred to by McGregor, and is
satisfied that the clouding described by McGregor is similar to that which is
found in the Hartley rocks.

Excellent examples of the brown clouding have been observed in a section
of the Odenwald gabbro, and in a uralitized dolerite from Broken Hill (Browne,
1922). Other evidence shows that both these rocks have suffered only a low
grade metamorphism.

The brief descriptions given above together with the detailed petrography
of the Hartley rocks points fairly conclusively to there being a relation between
the size of the inclusions and the grade of metamorphism—the higher the grade
the larger the inclusions. It would seem, therefore, that ‘‘clearing-with-inclusions”
and “clouding’’ are essentially the same phenomenon, the fundamental difference
being the degree of metamorphism suffered by the felspar. It has been pointed
out that McGregor has observed this with regard to the grey and brown clouding
and it would seem that clearing accompanied by granules or schiller plates
belongs to the higher grades of thermal metamorphism, whilst the clear felspar
unaccompanied by inclusions occurs when the grade of metamorphism has been
high enough to cause complete recrystallization of the felspar. It is pertinent to
note that the cleared felspar with granular inclusion is contained in rocks that
bear other evidence of a high grade thermal metamorphism. There is often
field evidence for this assumption, as well as the petrographic evidence of a
partial granoblastic structure, a preponderance of pyroxenes over amphiboles
and a marked development of rhombic pyroxene (Tilley, 1923). Sometimes, as
in the Broken Hill and Hyre’s Peninsula dolerites, brown hornblende is the most
abundant femic mineral and this indicates a high grade thermal metamorphism
under ‘wet’ conditions.

When both clearing and clouding are present, as in the rocks from Eyre’s
Peninsula, the Hart’s Range and Hartley, there is some doubt as to whether the
rocks represent an intermediate grade of metamorphism or whether one grade
has been superimposed upon another. From a consideration of the other evidence
it seems certain that at Hartley, at all events, the weaker grade has followed
upon an earlier high grade of metamorphism. The field evidence and the micro-
scope examination point to an earlier partial recrystallization of some of the
Hartley gabbros, followed by either reaction or hydrothermal metamorphism under
low grade conditions. With regard to the Hart’s Range rocks, the present writer
knows nothing of their field-occurrence, but the presence of green hornblende and
uralite wrapping partly recrystallized felspar crystals is certainly not contrary
to the assumption that a high grade metamorphism was followed by one of
lower grade. In the two Hyre’s Peninsula slides examined, there is no evidence
for assuming two grades of metamorphism, but Dr. Tilley himself states that
the rocks have suffered a high grade thermal metamorphism and other changes
during its decline (Tilley, 1921, pp. 117, 121, 125). The writer would suggest
that the slight clouding of the felspar might be attributed to this latter period.

BY GERMAINE A. JOPLIN. 155

With regard to a discussion on the origin of these phenomena, many difficulties
present themselves. It is obvious that iron oxides and/or magnesia are necessary
for the formation of the inclusions, whether they be ultramicroscopic or compara-
tively large as in the Broken Hill dolerites, and that the amount of available
material, besides the grade of metamorphism, possibly influences the size of the
inclusions. It would also seem that magnesia enters into their composition only
under the higher grade conditions.

Four possible explanations of the origin of the inclusions suggest themselves:
(1) Oxides held in solid solution in the original felspar. (2) Breaking down of
original inclusions in the felspar. (3) Addition of material from some external
source. (4) Previous alteration of the felspar.

McGregor has discussed this question and concludes that the iron and/or
Inagnesia are constituents of the original felspar. He states that the clouding
is confined to the more basic zones of the felspar, and that analyses of plagioclases
show that small quantities of these oxides are present in the more calcie end-
members of the series. In the felspar of the Hartley rocks, however, clouding
quite frequently occupies the more acid zones, and may be confined to certain
twin lamellae of the felspar, so it is evident that McGregor’s conclusions on this
point are not fully justified. Moreover, little reliance can be placed on chemical
analyses of plagioclases when such small amounts of “foreign” oxides are
involved, as it is impossible to free the felspar from original minute inclusions.

The schiller plates in the Hart’s Range and Dundas rocks certainly suggest
the squeezing out of material held in solid solution, but the amount of material
held in this way must necessarily be small, and it is difficult to account for the
numerous, comparatively large granules in the felspar of the Broken Hill dolerites.
Again, if clouding be superimposed upon granules, as suggested above, it is
difficult to account for sufficient material for the formation of both, since there
appears to be no diminution of either when the two phenomena are present
together.

If the material be derived from original inclusions, it would seem certain
that, under the low grade conditions, instead of clouding, a preliminary breaking
down of the original inclusions would be noticed.

That the material is derived from an external source seems out of the
question, since, in those Hartley types that have suffered the greatest deuteric
or hydrothermal alteration, there is little or no clouding present. Besides this,
it is evident that the hydrothermal solutions have produced alteration phenomena
of a different type—saussuritization, epidotization, etc. In accounting for schiller
inclusions, Judd (1885) ascribed them to percolating solutions under very deep-
seated conditions.

McGregor has considered the possibility of an earlier alteration of the
felspar, and his Ayrshire work (1930) has led him to the conclusion that only
those parts of the felspar that were originally fresh are now clouded. The present
writer feels that the same can not be said of the Hartley felspars. It is obvious
that most of the Hartley types (Joplin, 1931) have suffered some deuteric altera-
tion, and that the unmetamorphosed hornblende-gabbro in the centre of the Cox’s
River intrusion has been similarly affected; thus it would be too much to assume
that the rocks now represented by the recrystallized pyroxene-gabbros have not
also suffered from previous deuteric activity.

156 PETROLOGY OF THE HARTLEY DISTRICT, li,

It might be pointed out in conclusion that all the examples exhibiting these
phenomena, that have been discussed above, are basic rocks, and such types are
very prone to deuteric alteration.

SUMMARY AND CONCLUSIONS.

It has been shown that an earlier partial magma of gabbro differentiated,
giving rise to a slightly more acid core. The whole was then enveloped by a
ring-like intrusion of a later, more acid, partial magma of quartz-mica-diorite,
and the gabbros have thereby suffered three types of metamorphism: (1) Thermal
metamorphism. (2) Reaction or partial hybridization. (3) Hydrothermal
metamorphism.

It has also been shown that the gabbro and quartz-mica-diorite magmas are
consanguineous, and that the rock types arising from the reaction of the one
upon the other readily find a place upon a variation-diagram which is typical of a
normal calcic series. The close connection between hybridization and the internal
migration of magmas has been commented upon (p. 142), and it is assumed that
the Hartley rocks are hybrids, since the recrystallization of the gabbros evidences
an appreciable rise of temperature, whilst reaction, in the normal course of
differentiation, necessitates no such change.

It has been shown, therefore, that assimilation, as well as gravity separation,
has been an important factor in giving rise to the Hartley series.

On account of the close resemblance of the normal differentiates and the
hybrids arising from cognate parents, the writer concludes that a careful search
will reveal evidences of similar hybridization in other apparently normal rock
series—particularly among the basic members of the series. If such can be
shown, a strong case is made for assimilation as an important factor of differen-
tiation. At Hartiey the recrystallized-gabbros gave the clue to this interpretation,
and it has been possible to trace all gradations from one type into another. When,
however, the earlier intrusion is represented only by a very small outcrop, most
of the evidence may be obliterated.

In the present paper it has been suggested that the tiny granules of
birefringent minerals and iron ores in the cleared felspar indicate a high grade
thermal metamorphism, and the writer now makes the suggestion that these may
prove useful, as they have at Hartley, in revealing evidence of an earlier recrystal-
lization and so showing the rock to be a hybrid.

The origin of peculiar ellipsoidal bodies within the recrystallized rocks has
been discussed and chemical evidence adduced in support of the suggestion that
they represent recrystallized zoned hornblende crystals. At the same time the
possibility of contamination followed by recrystallization has not been entirely
dismissed.

It has also been shown that another rock of mixed origin has arisen as a
result of the contamination of the gabbro magma by cale-silicate sediments.

Acknowledgments.

In conclusion, the writer wishes to thank Professor L. A. Cotton, who has
made the time available for the completion of this work; Dr. Ida A. Brown
for kindly reading the manuscript, and Miss F. M. Quodling for assistance with
some of the optical tests. She should especially like to express her gratitude to
Assistant-Professor W. R. Browne for much helpful advice and the many valuable
Suggestions that he has offered during the preparation of this paper.

Proc, Linn. Soc. N.S.W., 1933.

GEOLOGICAL SKETCH MAP

OF THE

COX’S RIVER INTRUSION

a a

fT SS See

a
=

LEGEND

~~ |Alluvium

“+ | Contact-Allered
Upper Devonian Rocks

Eee
Quartz-mica-diorite
Ge

‘Diorlte- Gabbro and
Homblende-Gabbro

Hydrothermally Altered
| | Pvroxene ~ Gabbro

Hydrothermally Altered
" Reaction"= Gabbro

"Reaction" or

Hornblende - pyroxene Gabbro

Recrystallized
Pyrosene-Gabbro

Deuterically. Aliered
Homblende-Gubbro

9

Hedenbergite Ellipsoidal Bodies

7

aie ee ae Sree ceeg ls

Proc, Linn. Soc, N.S.W., 1933. PLATE VI.

val
ies

t
‘

Ne . i
a. c
wiRStes Benoa EP vs:
‘ ~
'
Me
ss
i
= \
,
y
.

BY GERMAINE A. JOPLIN. 157

For hospitality and help during fieldwork, the writer records her indebtedness
and thanks to Mr. and Mrs. J. Commens of Duddawarra, Little Hartley. She also
wishes to thank Dr. G. D. Osborne for his kindness in communicating the paper
to the Society during her absence from Sydney.

EXPLANATION OF PLATES V-VI.

Plate v.
Geological sketch map of the Cox’s River intrusion.

Plate vi.

1.—Deuterically altered hornblende-gabbro showing sub-ophitic fabric, and cores
of colourless augite surrounded by brown hornblende. Ordinary light. x 13.

2.—Recrystallized pyroxene-gabbro showing partly granoblastic pyroxene (augite
and hypersthene) and cleared felspar with tiny granular inclusions. Ordinary light. x 13.

3.—Hydrothermally altered pyroxene-gabbro showing uralitized partly recrystallized
pyroxene, plagioclase showing slight clouding superimposed upon previously formed minute
granular inclusions, and an amphibole vein cutting the rock. Ordinary light. x 13.

4.—Hornblende-pyroxene-gabbro showing “reaction’’ hornblende enclosing cores of
pyroxene, and a highly poikilitic relation towards partly dissolved and corroded crystals
of plagioclase. Ordinary light. x 13.

5.—Hornblende-gabbro showing brown hornblende wrapping plagioclase and suggestive
of the primary ophitic fabric (cf. Plate vi, fig. 1). Ordinary light. x 13.

6.—Felspar showing clearing and zonally arranged granular inclusions. Ordinary
lights x 23;

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Cc

158 PETROLOGY OF THE HARTLEY DISTRICT, ii-

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AUSTRALIAN COLEOPTERA. NOTES AND NEW SPECIES. VIII.
By H. J. Carrer, B.A., F.E.S.

(Hight Text-figures.)

[Read 31st May, 1933.]

Family BUuPRESTIDAE.
STIGMODERA (CASTIARINA) SUBTINCTA, Nl. SD.

Oblong-ovate. Head, pronotum, scutellum, appendages and the greater part
of underside golden green; elytra, save for the dark basal border, yellow with an
orange tint—the latter especially around the apex; the apical four segments of
abdomen yellow, sometimes showing faint metallic gleams.

Head deeply excavate and channelled, densely punctate. Prothorax: Apex
feebly, base more decidedly bisinuate, sides widely bulging at posterior third,
thence lightly converging to apex and more abruptly and subsinuately to base;
front angles acute, hind rectangular, medial line shown by a short sulcus near
apex and a smooth line on basal half; dise strongly, irregularly punctate, the
postero-lateral punctures coarse, contiguous and subrugose, elsewhere the punctures
becoming smaller, dense near apex, more distant towards base. Scutellum with
about six small punctures. Elytra widening at shoulders, lightly compressed
behind them, widest near middle, each strongly bidentate at apex, with long
external spine; margins entire, seriate punctures large and round in four sutural
rows, much smaller in the rest; intervals almost flat and impunctate throughout.
Flanks of pro- and metasternum coarsely punctate, the middle areas much more
finely so; abdomen glabrous with very fine shallow punctures. Dimensions:
15 x 5 mm.

Habitat.—Western Australia: Lake Grace (H. W. Brown).

This and the following species sent by that indefatigable collector, Mr. Horace
Brown, add to an increasing group having wholly or almost wholly yellow elytra—
Nos. 138-18 in my Revision (Aust. Zool., 1931, p. 350). The above species is
distinguished from all of these except intacta mihi, by its strong apical spines.
From intacta it is distinct by its smaller size, flatter form, and very different
prothorax. Holotype in Coll. Carter.

STIGMODERA (CASTIARINA) SUBNOTATA, N. Sp.

Oblong-ovate. Head, pronotum, scutellum, appendages and underside—except
abdomen—golden green (pronotum more golden than green), elytra and abdomen
testaceous, the former with a small black postmedial spot on the seventh interval.

Head excavate and channelled, finely, closely punctate. Prothorax: Apex
lightly, base more strongly bisinuate, sides moderately and nearly evenly rounded,
widest at middle, all angles rather wide; discal punctures fine and dense on
apical half, larger and more distant towards base, coarse near hind angles;
medial line only indicated by smooth area near base. Elytra lightly enlarged

160 AUSTRALIAN COLEOPTERA. NOTES AND NEW SPECIES, Vili,

near middle, apices finely bidentate with small circular excision; margins entire,
seriate punctures large and close; intervals finely punctate and nearly flat except
at base and extreme apex. Underside glabrous, sternal regions finely and closely
punctate, abdomen with dense shallow punctures. Dimensions: 11-12 x 4 mm.

Habitat—Western Australia: Dedari (H. W. Brown).

Three examples sent (1 g, 2 2) are similarly coloured, with the two small
spots varying slightly in size. It can thus be readily separated from Nos. 13-18
of my Revision or, if included amongst the species having spotted elytra, Nos.
39-53, it is entirely unlike any of these. Holotype in Coll. Carter.

STIGMODERA (CASTIARINA) INUSITATA, 1. Sp. Text-fig. 3.

Oblong, robust. Head and prothorax dark bronze, elytra red, with basal
border, two fasciae and apical mark blue-black, abdomen yellow, the rest of
underside black, mouth and legs blue, antennae green.

Head excavated and channelled; closely punctate. Prothorax widest near
base, apex arcuate-emarginate, anterior angles acutely produced; base lightly
bisinuate, posterior angles obtuse; sides rather gibbous at basal third, thence
obliquely narrowed to apex and base; disc closely punctate; punctures small
at middle, larger at base and sides, coarse near hind angles. EHlytra very slightly
widened at shoulders, scarcely compressed at middle; apices almost conjointly
rounded, a minute divergence only apparent; subapical margins very finely serrate;
striate-punctate, the strial punctures little evident; intervals very lightly convex
at base and apex, elsewhere flat, in general with transverse wrinkles and a
single row of large punctures. Underside with fine, close pubescence. Dimensions:
16-17 x 6 mm.

Habitat—wWestern Australia: Lake Grace (H. W. Brown).

Two examples sent, one evidently (4, are the only specimens I have ever seen,
and have been recently discovered by the alert enthusiast who has done such wide
entomological exploration in Australia. The form of the prothorax is much like
that of S. robusta Saund.; the elytral fasciae wide as in S. thomsoni Saund.,
but their colour is more nearly black, the premedial fascia not extending to the
sides; the apical mark is as in S. audax Saund., a broad, subapical arrow,
produced in the middle to cover the apex. Holotype in Coll. Carter, the second
example returned to Mr. Brown.

The following two recent captures of Mr. Horace Brown have just been
received (13/10/32) and show the almost endless tale of species in this genus:

STIGMODERA (CASTIARINA) RADIANS, n. sp. Text-fig. 1.

Oval. Head, pronotum and antennae bronzy green; elytra yellow on basal
half, red at sides and apical regions with blue markings as follows: wide post-
medial fascia extending to sides, lunate preapical mark, suture throughout, and
a wide basal band throwing off three branches, a sutural and two humeral, the
last covering the shoulders and extending obliquely backwards; underside and
legs dark green, moderately pubescent.

Head and pronotum closely, almost uniformly, punctate, the former channelled,
the latter with smooth medial line on basal half. Prothorax moderately convex,
apex and base bisinuate, the latter the more strongly; all angles acute; widest
behind middle, sides lightly, arcuately converging to the front. Scutellum sub-
cordate and concave. Elytra rather depressed and explanate, feebly obovate,
well widened at shoulders, apices with small, very briefly bispinose, lunation,
subapical margins almost entire (feebly serrulate as seen from beneath); striate-

BY H. J. CARTER. 161

punctate, strial punctures small, intervals strongly convex towards sides and
apex, lightly so at middle and clearly punctulate. Dimensions: 11-13 ~ 4-5 mm.
Habitat—Western Australia: Wembley (Mr. H. W. Brown).

E.H.Zeck Del.

Text-figs. 1-5.
1.—Stigmodera (Castiarina) radians. n. sp. 2.—Stigmodera (Castiarina) magnetica,
n. sp. 3.—Stigmodera (Castiarina) inusitata, n. sp. 4.—Microtragus discospinosus, N. SP.
5.—Saragus abnormis, n. sp.

162 AUSTRALIAN COLEOPTERA. NOTES AND NEW SPECIES, VIii,

Two examples sent by this tireless hunter of the genus suggest in general
form and coloration S. delectabilis Hope, or some examples of S. septemguttata
Waterh. var. tyrrhena Blkb.; the humeral extension somewhat as in SV. tantilla
Obenb., but shorter. In my Revision (Aust. Zool., 1931, p. 358) it would come
under group ix near anchoralis L. & G. Holotype and paratype in Coll. Carter.

STIGMODERA (CASTIARINA) MAGNETICA, 1. Sp. Text-fig. 2.

Oval. Head, pronotum and underside coppery bronze (in one example with
a suggestion of green), elytra testaceous with dark metallic-green bronze markings
as follows: basal band throwing off three branches, two short, irregular, juxta-
humeral marks parallel to but not extending to sides, a sutural branch expanding
behind scutellum into a diamond-shaped mark and joining the wide postmedial
fascia, this widest at suture and narrowing towards and meeting the sides; a
large triangular apical mark covering, or not quite reaching, the apex. Underside
rather strongly pubescent; legs dark green.

Head deeply excavate and channelled, strongly punctate, eyes unusually large
and prominent. Prothorax rather strongly convex, widest at middle, sides well
and evenly rounded, apex lightly, base more strongly bisinuate, anterior angles
acute, hind angles subobtuse; disc closely rugose-punctate, more finely so towards
middle, a smooth medial line traceable throughout, terminating near base in
an elongate coppery fovea. Scutellum coppery, subcordate. Elytra lightly enlarged
at shoulders, sides nearly straight; apices with a small, scarcely spinose, lunation;
margins clearly serrulate on apical third; striate-punctate, the strial punctures
small but distinct, intervals rather strongly convex on apical half, each with a
single line of impressed punctures. Dimensions: 12 x 44 mm.

Habitat.—Western Australia: Mount Magnet (Mr. H. W. Brown).

Two examples, the sexes, sent, that I name from its region, as also from its
dark metallic colour. The general style of pattern is not unlike that of the
preceding species, but the humeral mark leaves the shoulders uncovered, the
post-scutellary mark is wider as also are the fascia and apical mark. These last
three are connected along the suture. The apical mark is much as in S. audar
Saund. The colour of the elytral pattern is a very dark olive-green that at
first appears black, but in a good light its metallic gleams are evident. The short,
widely rounded and convex prothorax, with its rugose sculpture is distinctive.
Holotype ¢ and allotype ? in Coll. Carter.

STIGMODERA (CASTIARINA) BOOYANIA, Nl. SD.

Elongate-oblong. Head, thorax, underside and legs a rather brassy bronze
(pronotum greenish here and there), antennae greenish, elytra yellow with
violet-black markings as follows: wide basal margin, postmedial fascia (continuous
to sides), nearly straight apical mark, the suture (widely throughout and forming
an elongate diamond on basal two-thirds), and a short oblique, post-humeral
mark.

Head excavate and channelled, interior margins of eyes parallel, irregularly
punctate—closely at base, more sparsely on excavation, with laevigate area on
vertex. Prothorax convex (subgibbous), widest at middle, apex subtruncate
(slightly produced at angles), base lightly bisinuate, posterior angles produced
and acute; sides evenly rounded; disc with smooth medial line, the punctures
small, round and close on front half, larger near base, still larger and more
sparse at sides with some smooth rugae. Scutellum scutiform, strongly concave.
Elytra subangular near shoulder, sides subparallel (very lightly compressed near

BY H. J. CARTER. 163

middle), each apex with small, wide, shallow excision, margins subentire*; striate-
punctate, all intervals convex throughout, strongly and closely punctate. Sternal
regions with large, shallow punctures, abdomen densely and finely punctate.
Dimensions: 15 x 5 mm.

Habitat.—Western Australia: Booyana, Norseman district (Miss A. If. Baisiou).

A unique specimen, labelled as above, has been sent for my inspection by
Mr. F. EB. Wilson. I cannot find any species that approaches it very closely. The
figure of S. propinqua mihi (Trans. Roy. Soc, S. Aust., 1916, pl. 10, f. 19) is
rather near it, but this has a blue head, thorax and underside and the suture is
not dark throughout. It is much larger than S. bogania mihi, with a quite
different elytra and apical structure. The combination of subgibbous prothorax,
convex and strongly punctured elytral intervals, oblong form, colour and pattern
should render it clearly recognizable. The colour of the prothorax and its
punctures is somewhat as in S. undulata Don., though differently shaped. Holo-
type in Coll. Wilson.

STIGMODERA (CASTIARINA) HILARIS Hope, var. INFASCIATA, D. var.

The freakish variations of the genus are again shown here. Two specimens
taken by Mr. E. Sutton at Fletcher, S. Queensland, apparently belonged to an
undescribed species, but a further consignment of seven included one that was
identical with the typical hilaris. The others differ in the absence of the post-
medial fascia, the scutellary green marking and, in some cases, of the preapical
mark, with a prevailing orange ground colour. S. hilaris is not uncommon from
Bulli to Victoria; but hitherto I have not seen examples from regions north of
Sydney or variations from the typical pattern. It is thus desirable to give this
variety a name. It may be considered as a local subspecies, but this term has
been so much misused that I prefer to use the term variety.

CISSEIS SAPPHIRA, N. SD.

The whole surface, including appendages, peacock blue (in one example of
four, greenish), elytra with two indistinct pubescent fasciae, one extending in an
irregular zig-zag across the full width at apical third, the other, very short,
preapical.

Head channelled, scarcely excavate; densely and finely punctate. Prothorax:
Apex and base strongly bisinuate, widest near base, sides arcuately narrowed to
apex, all angles obtuse, the anterior depressed; upper carina continuous to apex,
the lower just visible from above; disc with fine transverse strigae, especially
near base and hind angles; two elongate depressions symmetrically placed on
basal half. Hlytra: Shoulders rather tumid, widest behind middle, subapical
margins finely serrulate, surface with fine silky scales; underside glabrous, nitid
and finely punctate. Dimensions: 5-6 x 24 mm.

Habitat.—Western Australia: Moore River (H. W. Brown).

Four examples examined show a short, rather wide species that, by colour,
can only be confused with (a) pulchella Cart. from N. Queensland, and (bd)
westwoodi L. & G. from Victoria and New South Wales, but (a) is larger, with
brilliant coppery head and prothorax and sides of prothorax nearly straight,
(0) is narrower and more elongate, golden green in colour with differently placed
pubescence. Type series in Coll. Carter.

* The faint crenulations, discernible through a binocular, are merely the result of
the coarse punctures of the intervals, not serrations as seen, for example, in S. cupricollis
Saund.

164 AUSTRALIAN COLEOPTERA. NOTES AND NEW SPECIES, Vill,

PARACEPHALA BROWNI, Nl. Sp.

Subcylindric. Head and thorax dull black with metallic gleams. Elytra
subnitid blue (sometimes purplish); underside and appendages bronze. Upper
surface with sparse, recumbent white hair, more evident on head.

Head convex, clypeus triangularly excised, forehead with fine medial sulcus.
Prothorax: Apex subtruncate, base strongly bisinuate, sides subparallel, anterior
angles depressed and rounded off, posterior subrectangular. Disc densely rugose-
punctate, a sinuous transverse depression across middle. Elytra finely granulose-
punctate, with some faint subobsolete costae, apices separately rounded and finely
serrulate. Dimensions: 5-7 x 13-2 mm.

Habitat—Western Australia: Merredin (H. W. Brown).

Five examples sent show a species allied to P. cyaneipennis Blkb., but
distinguished by the more sombre colour of head and thorax, the different blue of
the elytra and its different sculpture, with a finer silky surface. Blackburn’s
species, moreover, shows no sign of costae. Type-series in Coll. Carter.

Family TENEBRIONIDAE.
COTULADES SQUAMOSUS, DN. SD.

Brown, opaque, thickly clad with reddish squamose derm. Underside dark
brown, with evenly spaced setiferous punctures, each bearing a short adpressed
red hair.

Head rounded in front, covered with coarse recumbent scale-like hairs; eyes
coarsely faceted, antennal orbits raised, antennae very wide, 1-3 equally wide,
larger and longer than the succeeding, 4-8 gradually and slightly diminishing in
width, 9-10 wider than 8, sharply angulate at apices, 11 considerably narrower
than 10 and round. Prothorax wider than head, convex, apex subtruncate, sides
widest at middle, thence sinuately converging behind and arcuately to the apex,
crenulated by lateral scales, all angles obtuse, base rounded and slightly produced
backwards in the middle; surface uneven, discal area raised in front and behind,
and by more or less parallel ridges on each side of, a central depression, this
almost devoid of scales. Elytra obovate, subdepressed; at its widest much wider
than prothorax, with the suture raised and each with three wide subcostate
ridges, each interval between ridges with about three large depressions similar
to that on pronotum. Dimensions: 44 x 2 mm.

Habitat.—Victoria: Otway Brush Forest (F. E. Wilson).

Two examples taken by Mr. Wilson. A. wide, densely squamose species,
nearest to C. tuberculatus Cart., but differing in clothing, less prominent tubercles
and flatter form. Holotype in Coll. Wilson.

COTULADES TENUIS, DN. Sp.

Narrowly ovate, convex, rusty brown; the darker ground surface sparsely
clothed with brown derm; appendages paler; the unabraded part of surface
showing fascicles.

Head rounded in front, sides gradually converging behind, thickly, not
contiguously, punctate; antennae relatively longer than in OC. fascicularis,
segments 1-3 longer than the rest, the first widest, 4-8 subequal, 9-10 wider than
preceding, strongly transverse, 11 narrower than 10. Prothorax convex, oblong-
ovate, slightly longer than wide, produced in middle at apex and base, front
angles wide but recurved, hind angles obsolete, surface closely and coarsely
punctate, some white fascicles near apex and sides. Scutellum not visible. Elytra
closely fitting and of same width as prothorax at base, soon widening behind

BY H. J. CARTER. 165

this, rather narrowly tapering to apex; three little raised costae traceable on
basal half of each, intervals foveate-punctate with some seriate arrangement,
but largely hidden by derm; a few individual fascicles standing upright at sides
and apex. Legs with uneven surface due also to fascicles. Dimensions: 4 % 1-5 mm,

Habitat.—Victoria: Warburton District (Mr. F. E, Wilson).

A species distinct from the three recorded, not only by its different sculpture
but especially by the narrower form of both prothorax and elytra. Holotype in
Coll. Wilson.

(?) CoruLADES ABNORMIS, Nl. Sp.

Narrowly ovate; subnitid black, somewhat obscured in parts by brown derm,
upper surface with sparse long upright yellow hair, chiefly on elytra, legs brown.

Head subquadrate, truncate in front, canthus well raised, eyes large and
coarsely faceted, surface uneven through the presence of tubercles. Antennae
coarsely submoniliform, the two basal segments larger than rest, 3-8 subequal,
9-10 wider than preceding and subconic, 11 smallest. Prothorax convex, about
as wide as long, apex subtruncate, base rounded—produced in middle—widest
at apex thence lightly narrowing to base. Surface longitudinally corrugated by
lines of tubercles, of which the middle three form continuous costae. Scutellum
just perceptible. Elytra wider than prothorax at base, regularly ovoid, widest
at middle; surface sulcate-punctate, the punctures (where discernible) subfoveate,
more or less filled with flocculence, the raised intervals nitid, more or less uniform
at middle, at sides broken up into tubercles. Legs subglabrous and _ nitid.
Dimensions: 4 x 1:5 mm.

Habitat.—South Australia: Adelaide (Hart, 1886) in British Museum.

A single example sent by Mr. K. G. Blair is labelled as above. While differing
from its congeners in its longer antennae, more coarsely faceted eyes, and its
distinct sculpture, it presents so many affinities with the former species that for
the present it may be classified as above. Holotype in the British Museum.

COTULADES ALPICOLA, N. Sp.

Brownish-black; antennae brown, upper surface sparsely clothed with red
bristly hair, subrecumbent on head and pronotum, in part erect on elytra.

Head wide, truncate in front, depressed on vertex, rugose-punctate, antennal
segments 1, 2 very stout, 3 slightly narrower than preceding but longer than 4,
4-8 subequal, 9-10 trapezoidal, clearly wider than 8, 11 subspherical, narrower
than 10. Prothorax as wide as head at its widest, subtruncate at apex and base,
but the anterior angles clearly, though briefly, advanced; sides parallel and
coarsely crenulate; surface asperate and rugose. Elytra wider than prothorax
at base, ovate, relatively narrower than C. leuwcospila Hope; striate-punctate, the
suture and three rows of tubercles raised above the general surface, the inter-
vening space containing rows of subfoveate punctures, with small tubercles and
coarse setigerous clothing. Underside unevenly rugose-punctate. Dimensions:
3 SK IL saahoal,

Habitat—New South Wales: Mt. Kosciusko, 5,000 ft. alt. (the late Dr. E. W.
Ferguson).

Of a similar form to, but even smaller than, C. tenwis, but clearly separated by
coarser clothing, parallel-sided prothorax, and wider antennae. C. leucospila Hope
(= fascicularis Pase.) is larger, with white fascicles forming a pattern on the
pronotum, and without the large seriate punctures on elytra. Holotype in the
Ferguson Coll., Museum of Division of Economic Entomology, Canberra.

166 AUSTRALIAN COLEOPTERA. NOTES AND NEW SPECIES, Vili,

C. pilosus Oke, recently published with a brief description, is apparently
distinct from the above by (1) black setae of upper surface, also with certain
white setae, including two white spots on elytra, (2) antennae with “third
segment widest’. The white marks are apt to get obliterated in specimens of
the genus, but the striking difference of antennal structure seems decisive.

C. leucospila Hope = C. fascicularis Pase.—I am indebted to Mr. K. G. Blair
for this information, after a comparison of the types. This synonymy has not,
I think, been previously recorded.

The seven species of Cotulades, excluding pilosus Oke, which I have not
seen, may be tabulated as follows:

1 SIGeSNOfmprothora xa quitemonrencanlyaestralcihituyss)neeiiileeiicin eine eens 2
Sides of prothoraxs: widely, varcuacememicnaririciiac ne corinne ener nein eon aeRO 5
2. Antennae with 2 basal segments larger than rest ............... abnormis, Nf. sp.
Antennae iwith 3 basal segments! larzerithan’ rest {2c oss 2sen 2 cle stesso) cle lee 3
3. Antennal ‘segments; moniliform and marnower «.....--0.- see see eee: tenuis, n. Sp.
PN aUssoh WY eyewoaerdss> Wea eiss Enel WHE sagoogenogoubdsbbueGanoao0b0oboDb4O0 4
4 Hlyitray with jareeseriate: pun eckuresmmeieici sieiccia4 caeieitcaicienensls careiemeere alpicola, n. sp.
Bly trasowilth Out "Suh ey A Sey Oe Re Ae PRUNE chi avtalrauiatre Polke arta eviaentens Messin Ana ha lewcospila Hope
5. Pronotum and apical declivity of elytra coarsely tuberculate .... twberculatus Cart.
PronotumvandyapicaledeclhivitysoteelytrapnotmsOzseem oo ee eee eee 6
6. Elytral sculpture hidden beneath squamose derm .............. Squamosus, N. SP.
Hivtralgserlave punclunesmaiStin Cierieeieraciriel seme nacreren cai een montanus Blkb.

Platydema heroni Cart., described from Dorrigo (These Proc., 1929, p. 73),
has-been sent me since then by Mr. J. Armstrong, Bogan River, found in old
bird’s nest; also from Mr. Sutton, Fletcher, S. Queensland, in a similar habitat.

Mr. Jarvis (of the Queensland Department of Agriculture, Stanthorpe) has
found it in the nest boxes of finches in an aviary.

ENNEBOEUS GLABER, N. SD.

Elliptic; castaneous, nitid and glabrous, head and underside red-brown, the
former subfuscate; elytra a little clouded towards apex.

Head rather strongly punctate; antennae: 3-6 oval, 7 triangular, 8-10
increasingly transverse, cupuliform, 11 largest and round. Prothorax transverse,
widest at middle, apex and base lightly bisinuate, sides arcuate, anterior angles
rounded off, the posterior subrectangular; surface coarsely, irregularly punctate.
Scutellum oval, concave. Elytra closely fitting to and a little wider than prothorax,
everywhere rather thickly and coarsely punctate, without seriate arrangement,
the punctures larger than on pronotum. Legs short and stout, hind tarsi with
fourth segment equal to the first. Dimensions: 4 x 2 mm,

Habitat.—N. Queensland: Cairns, on sea beach (H. Hacker).

A single example in the Hacker Collection of the Berlin Museum clearly
differs from H. ovalis Waterh., and EH. australis Champ., in colour, absence of
pubescence, and antennae. In ovalis and australis segments 9, 10, 11 are said to
form a distinct but not large club, whereas in glaber 8-11 are clearly clavate. It
cannot be fossoris Oke, which has spinose tibiae and is doubtfully a member of
the genus. Holotype in Berlin Museum.

PTEROHELAEUS CASTANEUS, DN. SD.

Oval; nitid, dark castaneous above and below, foliate margins, antennae, palpi
and tarsi pale red.

Head rather flat, deeply sunk in prothorax, epistoma widely arcuate, its sides
little raised, surface densely, finely punctate. Antennae: segment 38 as long as
4-5 combined, 7-11 successively widened, the apical largest. Prothorax widest
near base, thence arcuately narrowed to front, and very slightly to the subacute

BY H. J. CARTER. 167

hind angles; apex arcuate-emarginate, anterior angles well rounded, base feebly
bisinuate, foliate margins wide and horizontal; dise with fine, shallow, scattered
punctures, the medial channel distinct, also two wide shallow basal impressions.
Scutellum arcuate-triangular, laevigate. Elytra as wide as prothorax at base,
widest near middle, foliate margins as wide as those of prothorax, subundulate,
and a little concave, narrowed at apex; seriate-punctate, scarcely striate; the
seriate punctures comparatively large, though varying in size, smaller in the two
sutural rows and in general so towards apex; intervals smooth and flat, a row
of larger punctures at junction with foliation; the fifth interval showing a
tendency to widening and feeble elevation. Prosternum sharply carinate, under-
side nitid and almost impunctate, abdomen finely striolate. Dimensions:
1} Se Mle} waabadl

Habitat—New South Wales: Mount Irvine (the author), Hornsby (C.
Gibbons).

I have long had a single example that would not fit any recorded description,
the colour suggesting immaturity; but the two specimens from Hornsby in the
Australian Museum are identical in colour, form and sculpture with this. I have
thus no longer any hesitation in describing it as a distinct species, nearest
FP. rubescens Cart., but smaller, with finer elytral sculpture; rubescens, moreover,
has the intervals raised. P. dispersus, of the black species, is somewhat like it
in form, but with smoother head and pronotum and larger seriate punctures of
the elytra, with irregular punctation of the basal area—quite absent from
castaneus. Holotype in Coll. Carter.

PTEROHELAEUS TENUICOSTIS, n. sp. Text-fig. 6.

Oblong, convex; opaque-black above, subnitid black beneath, legs brown,
antennae and tarsi rufescent.

Head concave, with reflexed margins, epistoma arcuate, surface thickly and
unevenly punctate; antennae with segment 3 shorter than 4-5 combined, 4-7
subconic, 8-10 rounded and transverse, 11 oval. Prothorax widest near base,
thence arcuately narrowed to front, and briefly so to base; apex widely, not
deeply emarginate, anterior angles widely rounded off and raised, base feebly
bisinuate, posterior angles acute and subfalcate, foliate margins wide, subconcave
and undifferentiated from disc by sculpture; disc towards sides subasperate with
shallow punctures; these tending to disappear near middle; in one example a
clear medial channel seen on anterior half, subobsolete in the second example.
Scutellum arcuate-triangular, its surface asperate. Elytra rather wider than
prothorax at base and thrice as long as it; subparallel for the greater part and
strongly convex; foliate margins narrow but of uniform width to apex; each
with seven very thin, sharp, little-raised costae, of which the first and seventh
are not very evident, the fourth and sixth with a slight tendency to crenulation
and towards apex to become nodulose; in the basal area each interval with an
extra costa—intervals between main costae with two rows of small punctures,
hese becoming indistinct towards sides; a row of large punctures at sides and
some irregular punctures in the scutellary region. Submentum and prosternum
rugose-pustulose; sides of mesosternum rugose, epipleurae pustulose, abdomen
punctate and striolate, legs coarsely punctate, the apices of tibiae and undersides
of tarsi strongly pubescent. Dimensions: 18 x 9 mm.

Habitat—Queensland: Coomooboolaroo Station (Rockhampton District) (J. R.
Slevin).

168 AUSTRALIAN COLEOPTERA. NOTES AND NEW SPECIES, Vili,

Two examples in the Australian Museum are very distinct from others of
Macleay’s Section ii, Subsection i. It is nearest, but not closely allied to,
P. elongatus Macl. The combination of oblong, convex form, opaque surface, and
finely costate elytra makes it easy to identify. Holotype and paratype (K 61089)
in Australian Museum.

SARAGUS ABNORMIS, n. sp. Text-fig. 5.

Elongate-ovate, rather flat, opaque brown; unabraded areas with bristly hair.

Head: Clypeus subtruncate in front, its sides meeting the canthus at a wide
angle; surface closely granulose, epistomal suture deeply impressed, terminating
in deep foveae; antennae strongly ciliate, segment 1 stout, 2 very small, 3 much
longer than 4; 5-8 successively shorter, 9-11 enlarged. Prothorax: Apex strongly
arcuate-emarginate, the acutely produced anterior angles almost concealing the
eyes; base bisinuate, much wider than apex; widest behind middle, sides lightly
converging behind, more strongly to the front, without sinuation; posterior angles -
subrectangular; margins with moderately wide foliation, a little obliquely raised,
without defined border; disc uniformly, densely granulose. Scutellum small.
Elytra of same width as prothorax at base, slightly widening behind middle,
humeral angle rather pronounced, a narrow horizontal margin extending a short
way beyond shoulder; surface irregularly costate as follows: a vague subundulate
costa on each side of and close to suture, more evidently raised at base and apex,
the suture itself costate towards apex; a second costa starting from base meeting
a third costa originating behind shoulder at the apical declivity, and thence
continued by a fragmentary costa not extending to apex; between the third costa
and margin a line of costulose tubercles; the interspaces between costae sub-
asperate, the extreme border—as with prothorax—subserrulate from bristly derm.

}

Text-figs. 6-8.
6.—Pterohelaeus tenwicostis, n. sp. 7.—Anaxo tenwicornis, n. sp.
8.—Homotrysis albolineatus, n. sp.

N. B. Adams, del.

BY H. J. CARTER. 169

Front tibiae
Dimensions:

Prosternum and epipleurae granulose, abdomen finely rugose.
spinulose on exterior margin, enlarged and bispinose at apex.
12 x 6 mm.

Habitat.—Mitchell (? Victoria River, N.T.).

A single example in the Australian Museum bears a label with the simple
word “Mitchell”. Mr. Musgrave informs me that this is probably in the vicinity
of the Victoria River, N.T. The form of the prothorax is somewhat like that of
Dysarchus and the elytra are flatter than usual with Saragus, but the eyes
are not divided, while the tibiae are without the characteristic spine of Dysarchus.
Type K 34307 in Australian Museum.

ONOSTERRHUS OBESUS, 0. SD.

Elongate-obovate. Nitid black above and below, antennae piceous red, its
basal segments and the tarsi red; tibiae and tarsi clothed with red tomentum.

Head: Labrum prominent, epistoma truncate, sinuate at sides making a wide
angle with the obliquely raised canthus; impunctate; antennae extending to basal
third of prothorax, segment 3 as long as 4-5 together, 3-8 more or less elongate,
9-10 round, 11 ovoid. Prothorax 5 x 8 mm., widest near middle, apex emarginate,
front angles rather wide and blunted at the tips, base feebly bisinuate, the dentate
posterior angles produced, twisted obliquely outwards; sides moderately rounded,
sinuate behind, the edges thickened, raised and channelled within; disc impunctate.
Elytra very convex, wider than the prothorax at base, about as long as wide
(11 mm.), widely rounded at shoulders, apical declivity steep, a narrow horizontal
margin, just visible from above, bearing a row of large punctures; dise very
minutely and sparsely punctate, each with three subobsolete costae traceable and
some vague unevenness and shallow wrinkles. Metasternum and abdomen with
longitudinal strigae, teeth of submentum bluntly enlarged and obliquely raised.
Dimensions: 19-20 x 10-11 mm.

Habitat: South Australia: Murray River (A. H. Elston).

Two examples have been in my cabinet for some years labelled O. stepheni var.,
but the following comparison will serve to distinguish these allied species:

O. obesus
Form: more convex and obovate
Surface: more nitid and uneven, elytral
costae traceable
Prothorax: narrower, its raised border
less thick
Submentum: teeth smaller and sloping
forward
Holotype in Coll. Carter.

O. stepheni Cart.
less convex and ovate
opaque and more even, no sign of
costae
wider, more rounded at sides, border
thicker
teeth larger and vertical

ONOSTERRHUS POLITUS, Nl. Sp.
Widely ovate; very nitid black above and below, antennae and tarsi reddish.
Head: Episterna concave in front, its margin well raised, almost rectangular
at the corners, at sides making a wide angle with the antennal sockets, these

rounded and raised;

forehead minutely punctate;
impression. Underside of head impunctate, tooth of submentum small.

with medial longitudinal
Prothorax

4 x 75 mm., apex arcuate-emarginate, its angles acutely produced and pointing
forward; base bisinuate, widest behind middle, sides sinuate in front and behind,
more widely so in front; lateral border very wide and convex, deflexed at hind

170 AUSTRALIAN COLEOPTERA. NOTES AND NEW SPECIES, Viii,

angles, channelled within; disc polished and impunctate. Scutellum strongly
transverse. Elytra rather wider than long (9 x 10 mm.), very convex, its narrow
horizontal margin only seen from above near shoulders; nitid and impunctate,
showing traces of subobsolete costae. Prosternum and sides of abdomen
longitudinally wrinkled. Posterior legs wanting. Dimensions: 15 x 10 mm.

Habitat.—North-West Australia (in National Museum).

A single example, slightly mutilated as to legs, is so distinct as to merit
description. In my table of the genus (Ann. Q’land Mus., 1911, p. 7) it would
stand next to O. rotundata Blkb., which is a larger species with muck less
prominent front angles to the pronotum. Holotype in the National Museum.

BYALLIUS OBERONIUS, N. SD.

Elongate-ovate; dull black above and beneath, oral organs and antennae
reddish-brown, tarsi and apex of tibiae with red tomentum.

Head densely and finely punctate, labrum prominent, epistoma truncate,
forming a right angle with the raised canthus, antennae having segment 3 as
long as 4-5 combined, 4-8 suecessively widened, 9-10 spheroidal, 11 ovoid. Pro-
thorax (4 x 6 mm.) widest near base, apex arcuate-emarginate, anterior angles
acute, produced obliquely outward, base wider than apex, truncate, posterior angles
obtuse, scarcely dentate, sides well rounded, with short sinuation behind and a
longer one towards the front angles, revolute border thick, round, and channeiled
within; surface finely and closely punctate with an obvious but fine medial impres-
sion on middle half, terminating behind in a light wide triangular impression;
this continued transversely parallel to base. Elytra ovate, narrower than
prothorax at base, shoulders roundly widened, widest near middle, surface very
uneven, irregularly costate-reticulate, each with three irregular costae, intervals
with well defined raised reticulation, depressed parts finely punctate; suture also
widely raised, extreme border unseen from above. Underside coarsely punctate,
the prosternum also transversely rugose, the abdomen longitudinally so except on
apical segment, this densely punctate. Dimensions: 16-18 x 7-8 mm.

Habitat—New South Wales: Hazelgrove, Oberon District (F. H. Taylor and
Consett Davis).

Seven examples before me show no external sexual distinction. It is nearest
to B. angustatus Cart. in elytral sculpture, and to B. ovensensis Cart. in its
rounded pronotum, but with the strongly thickened recurved border of the
former species. The elytral reticulations are more strongly defined than in
any other of the genus. Holotype in Coll. Carter.

The genus Byallius, not only by sculpture, but especially in the form of
the mentum, submentum, prosternum and long metasternum, is clearly more allied
tu Nyctozoilus than to Cardiothorax, near which the author placed it. The sub-
mentum is characterized by its wide, generally rough, surface, divided medially
at the base by a nitid triangular plate. The species are in some cases closely
allied and are very local. They may be separated into two groups by the elytral
sculpture:

Group A.—Species having defined raised reticulations, as in Nyctozoilus
reticulatus Bates.

Group B.—Species having the intervals between costae variously punctate,
rugose, or vermiculate, without closed reticulations.

B. reticulatus is unfortunately named since it comes clearly in Group B. Four
species have been added since my former table was published (These Proc., 1919,
Dp. 159), so that the folllowing amended tabulation is desirable:

BY H. J. CARTER. 171

Byallius.
(Mahvitralointervals with pronounced reticulations! 2.2. 022.1. -lelee so clels cles oles wieleislec se 2
Mivtralnintervals: without saehned meticulations: -)...... «++ <1 siecle « «ec cteee sicle ee 7
A CONOtAlOutluMmemILreLWlave (SUCHEN TILALC)) i crsisiene «coke ns: 6 eee el aieteleieneusieeleienel clelanele 3
IPrOnocalyOuclinementin emai oii sistece sl cicretseiolsiaichehet sileusie 2 eevee cueud Drevavenesetaveletsracpaxel oie eis 4

3. Pronotal border thick, raised, its hind angles obtuse without preceding sinuation
ROEM IRS tl cH Retires oe Bek one vtctrente dal oi anteyrortel tte teita| or. sicdicoNel esata), o's ch oi Shs,cé).e. VS“ eh ole to laticollis Cart.
Pronotal border less thick and raised, hind angles dentate with preceding sinuation

5 iON OCI STEN ONE UIE 6 BBD Gee PRO RTI CNET 0 OR ic eT ENT ROR 07 rem andersoni Cart.

APE CONOCUIM WAGE lym CONCAVerwACi IMD OLGEI meter cele ccneia tteleliel ciel ells) ella sisilae)clenaieiorehelete tale 5
Pronotum) withoutvappreciable’ concawity ]. 06 .cnsccias acces. kosciuskoanus Cart.
Dea SrOnoOLiuImmwitheGelimed medial Camaliver mics «o ccieievelelenenscl ssc 6 ce ce «6 oberonius n. sp.
CON OULU WALLED OUR CAT a lime wircnctet siren lets ciccsts) ciecer sltene rae ol omeueitey cleus ts <w1 6) clolaxe, olvoraiede “oneuenod severe 6

RT ea MN Pear Tiek ae oe cliente ustislichoutsie Tae fore iowol one) wivonsnshameloleae vale nave salle wie eve ovensensis Cart.

Sides of prothorax little rounded, front angles pointing forward .. angustatus Cart.
fAomeronotum subnitid® (elytral costae little raised’ Sia.c..cass+es 6. oe reticuwlatus Pasc.
PronotumPopagqueselytrallvcostae welll raiSed! sien. ie cis cue el cists cue sue sels se see cele 8
Ceuta tervals Oveace-pUmMCLates ysis: ai clerac che shehetel eee ie cicvele slael cra a cele mastersi Cart.
Ey brallaim terval Sit) Oteey SOwmysraiss oe deta sts deviclicire vorhouelonen Viste ekohoyioy o) shore) redone Len cnenetteurap ey Scenes ede: celTe\-eh o-foy 9
Seeeronotall border thick and well raised 3.02. ve ee cee re (Styrus) revolutus Cart.
Pronotalmborder Witter thickened (Or sraisSeG ejercicle clceice te ocla6 cele circle punctatus Cart.

Lyprops atronitens Fairm.—When examining some Tenebrionidae belonging
to the Australian Museum I was astonished to find an example of this species
which had been received from Mr. W. Heron, taken at Hast Dorrigo, New South
Wales. It is surprising that a common New Guinea species should occur so far
south in Australia, having hitherto escaped notice in the more northern regions
of the continent. There is no reason to doubt the evidence of this distribution.
This is the first Australian record of the genus (vide my note under Pseudolyprops
australiae Cart., These Proc., 1930, p. 538).

Chariotheca striato-punctata Macl. = C. doddi Cart. (var.)—Mr. Blair has
called my attention to the probability of this synonymy. After a close re-
examination of my type I must confess that it is too close for specific distinction
from the older Macleay species, though shorter and rather wider in form than
the examples before me of this.

MENEPHILUS ARMSTRONGI, 0. SDP.

Oblong; pitchy black, elytra reddish-brown subnitid, glabrous and more nitid
beneath; oral organs, antennae and tarsi red, tibiae and femora reddish in parts.

Head depressed, clypeus rounded in front, not raised at sides; coarsely and
closely punctate; antennae short, its four apical segments transverse. Prothorax:
Apex emarginate, anterior angles sharply produced (as in M. rectibasis Cart.),
base truncate, posterior angles obtuse and well defined; sides lightly rounded,
converging towards apex; a narrow horizontal margin, scarcely sulcate within,
disc strongly punctate, the punctures more distant than on head, becoming finer
at middle, a light elongate depression at base on each side near hind angles.
-Scutellum transversely oval, punctate. Elytra rather wider than prothorax, sub-
parallel, striate-punctate, the three striae near suture deeper than the rest and
containing smaller punctures, the outer striae shallow with larger punctures,
intervals wide and very little raised. Underside of head with strong, confluent
punctures, prosternum transversely rugose, its episterna finely pustulose; meta-
sternum and abdomen strongly, but not closely, punctate; anal segment clearly
margined. Femora tumid, hind femora with wide triangular enlargement near
apex. Dimensions: 8 x 24 mm.

172 AUSTRALIAN COLEOPTERA. NOTES AND NEW SPECIES, Viii,

Habitat—New South Wales: Nandewar Ranges (J. Armstrong).

Mr. Armstrong took two examples on our recent (November, 1932) trip to
this region, of which he generously gave me one. It is clearly distinguished from
M. humilis Er., by small size and glabrous sternum; from WM. colydioides Er.
(which it most resembles) by the less nitid surface, more finely punctate
pronotum, wider and less convex elytral intervals, besides colour differences.
M. rectibasis Cart., is easily distinguished by its very nitid black surface, the fine
sculpture and sulcate margins of pronotum, and the deep striae and sharp intervals
of elytra. Holotype in Coll. Carter. Paratype in Coll. Armstrong.

OLISTHAENA SPARSA, DN. SD.

Ovate, very convex; brilliantly nitid brown-bronze above and below, antennae
and tarsi red.

Head: Labrum prominent, clypeus subtruncate, eyes large, widely separated;
antennae rather short, segments 3-6 sublinear, 3 longest, 7-10 very little widened,
11 ovate, longer than 10; surface of head and pronotum smooth and impunctate.
Prothorax strongly transverse, broadly emarginate at apex, its angles subacute,
extending to half the eye, base strongly bisinuate, posterior angles acute and
faleate; widest at base, sides obliquely narrowed on basal half, thence arcuately
so to the front, a narrow raised border at sides enclosing the front angles, a
transverse depression near base, interrupted in middle, terminating in large
fovea near sides. Scutellum triangular. Elytra wider than prothorax, gibbous
in front of middle, a narrow lateral gutter throughout; surface with large, sparse
punctures, irregular in size and arrangement, those nearest suture smaller, else-
where foveate, but obsolescent on apical declivity. Underside as brilliant as the
upper and impunctate, save for a few large punctures on flanks of mesosternum.
Legs moderately long, hind tibiae a little flattened and widened in middle, post tarsi
with basal segment about as long as the apical. Dimensions: 11 x 6 mm.

Habitat— Queensland: McPherson Range (H. Tryon).

A unique specimen in the Queensland Museum is difficult to place, its slender
antennae and gibbous elytra distinguishing it from other members of the genus,
but it would be still more out of place in other genera of the subfamily. The
elytral sculpture is somewhat as in Prophanes mastersi Macl., on a smaller scale.
Holotype in Queensland Museum.

EUTOREUMA TURNERI, 0. Sp.

Hlongate-ovate, convex; nitid metallic greenish-bronze above, nitid black
beneath, legs black, antennae, tarsi and palpi red; basal segments of antennae
infuscate.

Head: Clypeus convex, forehead abruptly rising from the suture, eyes large,
rather close, interspace about the diameter of one eye; surface finely punctate,
antennae very slender, not extending to base of prothorax, 3-6 lineate, 7-10
submoniliform, slightly successively increasing in size, 11 largest, oval. Prothorax:
Apex deeply emarginate, the acute—but not sharply pointed—front angles
extending beyond the eyes; base bisinuate, sides arcuately narrowed from base
to front, with thin raised border and moderate foliation, disc with minute and
dense system of punctures, two small foveae near middle, two basal foveae, the
area between these also lightly depressed. Scutellum triangular. Elytra slightly
wider than prothorax at base, oblong, subparallel, convex, but scarcely gibbous,
lateral border not visible from above; surface unevenly embossed, the irregular
raised areas nitid and laevigate, the depressed areas with irregular clusters of

eo

BY H. J. CARTER. 17:

small, brilliantly coppery punctures, continued sparsely to the apex. Pro- and
mesosternum and abdomen striolate, metasternum sublaevigate; tibiae of ¢ rather
strongly curved and pubescent beneath, of 9 straight without pubescence.
Dimensions: fg, 12 x 5 mm., 9, 138 x 6 mm.

Habitat—Queensland: Eungella (Dr. A. J. Turner).

A pair—the sexes—of this fine insect is clearly congeneric with H. cupreum
by similarity of form, sculpture, and slender antennae. The brilliant punctures
with the more sombre embossed areas give a special character to the species,
which I gladly name after an old friend, its discoverer. Holotype and allotype
in the Queensland Museum.

ANDROSUS HACKERI, Nl. SDP.

Oblong-ovate; convex; nitid, head and prothorax black, elytra violaceous,
antennae and legs dark, the base of the former and the tarsi red.

Head and pronotum closely, not coarsely, punctate. Antennae gradually
widening from segment 5; 7-10 cupuliform, 11 round. Prothorax strongly trans-
verse, widest a little in front of middle, thence arcuately narrowed anteriorly,
and subrectilinearly to the base; apex and base lightly bisinuate, anterior angles
moderately produced and obtuse, the undentate posterior angles also rather wider
than 90°; a wide raised lateral border with a row of large punctures immediately
within it; front and hind margins without such border. Elytra convex, of nearly
same width as prothorax at base, slightly widening beyond middle; seriate-
punctate, the seriate punctures small and well separated in sutural rows, rather
larger towards sides, the series extending clearly to the extreme apex; intervals
flat. Prosternum densely and coarsely punctate, its process bisulecate with large
irregular punctures; episterna and epipleurae with large, sparse punctures;
abdomen with finer punctures, except near hind coxae. Dimensions: 6 x 24 mm.

Habitat—Cape York: Coen District (H. Hacker).

A single example in the Hacker Collection of the Berlin Museum is nearest
to A. brevis Cart. amongst Australian species, and to A. violaceus Pasec. amongst
New Guinea species, but differs from both in its black head and prothorax.
Holotype in the Berlin Museum.

ADELIUM FLAVITARSIS, 0. SD.

Rather widely ovate, depressed; head and pronotum very nitid black with
purple sheen; elytra brilliant metallic-purple in one example, dark violaceous or
cyaneous in the other two, prosternum, epipleurae and legs red, abdomen black;
palpi and tarsi yellow, antennae brown, with basal segments sometimes red.

Head sparsely but clearly punctate, forehead with two large foveae between
the eyes, antennae stout, not extending to base of prothorax, segment 3 shorter
than 4-5 combined, 3-6 moniliform, 7-10 cupuliform and successively widened,

1 largest, ovate. Prothorax transverse, widest at middle, apex arcuate, anterior
angles obtuse, base subtruncate, sides moderately rounded, subsinuate before
the defined but obtuse hind angles; extreme border narrowly raised throughout,
explanate margins defined, on anterior half by a short sulcus; dise indistinctly
and minutely punctate, a fine medial channel clearly impressed, not quite extending
to apex or base; a few ill-defined ridges, the most obvious being one obliquely
directed forwards from the posterior angles. Scutellum triangular. Elytra slightly
wider than prothorax at base and about twice as long as it; ovate, shoulders
well defined, a narrow border visible from above throughout; sulcate, the sulci
nearest suture more or less entire, the external sulci irregularly interrupted, in

D

174 AUSTRALIAN COLEOPTERA. . NOTES AND NEW SPECIES, VI1ll,

some cases forming elongate foveae with cancellate transverse ridges, without
perceptible seriate punctures; intervals convex, nitid and impunctate. Under-
side very nitid, vaguely punctate on prosternum and epipleurae; fore tibiae lightly
curved, post intercoxal plate truncate. Dimensions: 11-12 ~ 43-5 mm.

Habitat—Australia (Kraatz).

Three examples in the Berlin Museum sent for determination are very distinct
from all described species by brilliant colour, the suleate elytra and absence of
seriate punctures (A. striatum being the only other sulecate species). The elytral
sculpture is nearest that of A. ruptum Pasc. The difference of colour in the
specimens is not, I think, sexual, since all three are probably male. A. flavicornis
Cart. has similarly coloured tarsi, but is widely different otherwise. It is regret-
table that the attached labels only bear the words ‘‘Australien” and ‘“Kraatz’’. Holo-
type in the Berlin Museum.

SEIROTRANA SUTTONI, N. Sp.

Ovate; nitid dark bronze, tarsi red.

Head coarsely, irregularly punctate, forehead with a short medial sulcus
besides the usual stirrup-shaped impression. Antennae submoniliform, the two
penultimate segments transverse, the 11th largest, ovoid. Prothorax: Apex
arcuate-emarginate, base truncate, sides moderately rounded, widest at middle,
lightly sinuate behind, anterior angles rather strongly produced and acute,
posterior rectangular, not dentate; margins with a slight tendency to crenulation,
extreme border narrow; disc closely and irregularly punctate with a few rather
vaguely defined larger punctures. Elytra oval, wider than prothorax at base,
striate-punctate, seriate punctures small near suture, larger towards sides;
alternate intervals with inconspicuous nodules, except on the 7th and towards
apex; the 3rd and 5th only irregularly convex; about four large punctures on
the Ist and 3rd, and some minute pustules perceptible on the 4th and 6th.
Epipleurae punctate, prosternum transversely wrinkled, abdomen sublaevigate
save for the longitudinal strigae behind the post intercoxal plate (found in
several other species). Dimensions: 9-11 x 3:5-4:5 mm.

Habitat—Queensland: Stanthorpe (EH. Sutton).

Four examples differ from S. acuticollis mihi (also from Stanthorpe) widely
in the wider, narrowly bordered prothorax, coarser ground sculpture, and less
numerous and defined larger punctures. In common with S. anomala mihi, it
possesses characters of both Groups I and II of my revision, though most closely
allied to species of Group II. In form nearest, though less wide than, S. repanda
Pasc., its elytral sculpture is nearest minor mihi of Group I. Holotype in Coll.
Carter.

SEIROTRANA FUNEREA, DN. SD.

Oblong-ovate; opaque black above, nitid dark bronze beneath, antennae and
tarsi dark brown.

Head densely punctate, antennal segments 4-8 oblong conic, the two penul-
timate transverse and triangular, 11th large. Prothorax: Apex arcuate-
emarginate, base subtruncate (feebly concave), sides very lightly rounded, widest
before middle, sinuate and converging behind; anterior angles wide, posterior
defined but undentate and obtuse; margins finely crenulate, a narrow external
border, disc closely strigose-punctate, a large depression on each side, with
declivous lateral foliation having transverse strigae. Elytra convex, wider than
prothorax at base, shoulders rounded, a narrow raised border from above seen

BY H. J. CARTER. 175

only on basal half; striate-punctate, the seriate punctures small, the odd intervals,
including the sutural, with short studs (catenulations) regular in size, those on
the sutural small and opaque, with a puncture between each pair, those on the
érd, 5th, 7th, 9th larger and nitid; the even intervals bearing minute pustules.
Epipleurae with large, sparse punctures, prosternum coarsely, irregularly punctate,
abdomen sublaevigate. Dimensions: 11 x 4:5 mm.

Habitat.—Queensland National Park (R. Illidge).

A single example, sent by my old friend, is distinct from the other black
species of Group I. It is separated from NS. catenulatus Boisd., by smaller size,
much finer and closer sculpture of pronotum and less strongly crenulate margin;
from S. carbo mihi by its more opaque surface, the small seriate punctures of
elytra and generally impunctate intervals; and from both in having the small
catenulations on the sutural interval. Holotype in Coll. Carter.

SEIROTRANA MONTICOLA Blkb. var. STRIGIVENTRIS, N. var.

I have two examples of Blackburn’s species from Mt. St. Bernard (Victorian
Alps) that show the strong sexual characters described. Two other examples,
taken by myself at Mt. Kosciusko in 1906 and 1926 respectively, may be later
found to be distinct from monticola, but, having many characters in common, may
for the present be named var. strigiventris. The chief differences are as follows:

monticola Blkb. var. strigiventris.
Dimensions: 11:5 x 5 mm. wy Se) Gb Foahoay
Prothoraz. All angles more sharply defined.
Hlytral intervals 2, 4, 6, wide, 2 and 6 2, 4, 6 very narrow, 2 and 6 without
with a few large punctures. punctures.
Apical segment of abdomen “valde Lightly ridged, rather than carinate,
carinato”’, apex not excised. apex with triangular excision.

In both typical form and the variety the two basal segments of abdomen are
deeply longitudinally strigose in the middle. The crenulation of the pronotal
margins is subobsolete but traceable.

Note.——Seirotrana repanda Pase. = 8S. johnstonensis Cart.—The latter name
must be sunk. I was deceived by the distortion of my example, which a re-
examination makes clear.

DYSTALICA MACKAYENSIS, Nl. SD.

Oval; head and pronotum opaque black, elytra and underside subnitid red,
the former the darker; antennae fuscous; surface with short bristly hair.

Head densely and strongly punctate with granulose intervals, clypeus bilobed,
with a deep medial bay, antennal orbits widely concave, subangulate in front
of eyes;. vertex flat, antennae slender, 2-3 lineate, 3 as long as 4-5 combined,
4-7 subconic, 8-10 successively widened and rounded, 11 largest. Prothorax
widely transverse, widest near base, apex arcuate-emarginate, anterior angles less
than 90°, sharply defined; base strongly bisinuate, produced backward near angles;
sides well rounded, sinuate before the acute, but not dentate, posterior angles,
margins with slightly hollowed foliation not differentiated from disc by sculpture,
extreme border thin and entire, though outlined by the bristly hair; dise with
faint indication of medial impression, otherwise uniformly punctate-pustulose as
on head, the round punctures rather larger than on head. Scutellum transversely
triangular. Elytra slightly wider than prothorax at base and three times as long,
sides feebly widened behind middle, apex rather widely rounded; striate-punctate,

176 AUSTRALIAN COLEOPTERA. NOTES AND NEW SPECIES, Viii,

the rather large seriate punctures separated by cancellate ridges; intervals
steeply convex, the second and sixth more prominent than the rest, and trans-
versely rugose and finely granulose. Underside finely rugose, tibiae slender.
Dimensions: 8 x 4 mm,

Habitat.—Queensland: Mackay (W. A. Macdougall).

Holotype (unique) in the Queensland Museum.

Family CISTELIDAE.
ANAXO TENUICORNIS, nN. sp. Text-fig. 7.

Elongate, cylindric; nitid brownish or bronzy black, upper surface with a
short pale pubescence; underside nitid black and glabrous, basal half of femora,
also tibia (mostly) yellow; palpi, tarsi and remainder of legs black.

Head closely, finely punctate, rugose on the depressed rounded epistoma, eyes
widely separated, not prominent; antennae very thin, 5-10 subequal in length,
very lightly successively widening, 11 smaller and narrower than 10. Prothorax
subquadrate, transversely convex, aS wide as long, apex and base truncate, anterior
angles depressed and rounded off, the posterior obtuse, sides nearly straight;
disc with fine, close, shallow punctures; a large foveate depression at base, and
(in one example) two foveae near middle, without a sign of medial line. Scutellum
transverse. Elytra considerably wider than and thrice as long as prothorax,
shoulders squarely rounded, sides parallel for the greater part, separately rounded
at apex; striate-punctate, the striae fine, the punctures small and irregular,
intervals—at least the three nearest suture—convex and finely rugulose. Femora
moderately enlarged, tibiae lightly enlarged at apex. Dimensions: 9-10 x 24 mm.

Habitat.—New South Wales: Wheogo, 12 miles N.E. of Dunedoo (A. Musgrave).

Two examples taken by Mr. Musgrave are clearly distinct from A. cylindricus
Germ., by the much finer sculpture and fragile form, while the antennae are
finer than in any of its congeners. One example, with mutilated antenna, shows
a slight thickening of tibiae that suggests sexual distinction, but the fragile
nature of the specimens prohibits manipulation. Holotype in Australian Museum.

HoMOTRYSIS ALBOLINEATA, nh. Sp. Text-fig. 8.

Elongate-obovate; bronzy chocolate, subnitid above, very nitid beneath, whole
surface irregularly clothed with white recumbent pile.

Head: Clypeus subtruncate, front margin raised; eyes widely separated (as
in H. cisteloides Newm.), antennae finely lineate, segments 3-11 subequal. Pro-
thorax subconic, widest at base, subsinuate before the acute hind angles, thence
converging to and arcuate towards apex; disc closely and a little irregularly
punctate; medial channel distinct nearly throughout in one example (on apical
half only in the 2nd). Scutellum densely clothed with white pubescence. Elytra
considerably wider than prothorax at base, widest behind middle; striate-punctate
with a short scutellary and nine other rows of small punctures placed in fine,
clearly-cut striae; the seriate punctures very small except at sides and humeral
region; intervals flat, those from the fifth outwards densely pubescent, forming
distinct lines, especially when viewed from behind, the denuded intervals showing
fine transverse rugae. Legs densely, the underside sparsely, pubescent.
Dimensions: 13-14 x 6 mm.

Habitat—New South Wales: Wheogo, N.E. of Dunedoo (A. Musgrave).

Two examples, both, I think, ¢, are nearest to H. cisteloides Newm., and
H. scutellaris mihi—in size and form to the former, with the white scutellum of
the latter. From both it is readily distinguished by the fine seriate punctures,

BY H. J. CARTER. 177

the flat interstices and less strong sculpture of the elytra; besides the well-defined
linear pubescence on the external half. Holotype in the Australian Museum.

Homorrysis INTERSTITIALIS, MN. SD.

Oblong-ovate; nitid black, antennae, palpi, tarsi red, tibiae piceous red; upper
surface sparsely and lightly pubescent.

Head rounded in front, densely and finely punctate, eyes in ¢ closer than in ?
—in the latter the interspace about the diameter of an eye; antennal segments
subtriangular, 4-10 subequal, 3 longer than 4, 11 shorter than 10. Prothorax
subquadrate, slightly widest in front of middle, apex lightly convex, base bisinuate,
with wide medial lobe; sides nearly straight, anterior angles widely rounded and
depressed, posterior subrectangular; surface rather convex, lateral border unseen
from above, disc densely, subcontiguously punctate, in one example (?) showing
a depressed medial line. Elytra obovate, slightly wider than prothorax at base,
widest near apical declivity; striate-punctate, the striae very fine but distinct,
containing minute punctures, intervals flat and wide, and crowded with punctures;
these on basal half coarse, and more or less in geminate rows, becoming smaller
and more irregular towards apex, a sparse pubescence visible on sides. Pro-
and metasternum strongly punctate, the punctures more distant on the episterna.
Dimensions: 9-10 x 3-4 mm.

Habitat.—Brisbane (L. Wassell).

Four examples, two of each sex, show a species distinct through the form of
its prothorax and its unusual elytral sculpture; the former exceptionally robust,
the latter having its interstitial punctures greatly exceeding those in the striae
in size and prominence. 4H. laticollis Boh., perhaps its nearest ally, has the
prothorax widest at, or near, base, the surface punctures fine and distant, the
elytra having its strial punctures larger than the interstitial. Holotype ¢ and
allotype ¢ in Coll. Carter.

HoMoTRYSIS RUFO-BRUNNEA, TD. SD.

Elongate-ovate; nitid reddish-brown, oral organs, antennae and legs red;
sparsely pubescent.

Head and pronotum densely punctate, the punctures large on basal. region
of the latter; eyes widely separated, interspace equal to the transverse diameter
of an eye; antennae: segment 8 linear, 4-10 subequal, scarcely enlarged at apices,
11. more slender and pointed. Prothorax widest at base, thence lightly and
arcuately narrowed to apex, this slightly produced in middle; front angles
depressed and subobsolete, hind angles acute but blunted at extreme tips; base
bisinuate with light medial depression and two wide, shallow excisions at base.
Elytra wider than prothorax at base and slightly obovate, striate-punctate,
intervals flat, the striae sometimes obliterated by the coarse punctures of the
intervals, irregularly scattered but considerably larger than the indistinct small
punctures in striae, and increasing in size laterally; sides and apical region
with rather long subrecumbent pale hair. Metasternum sparsely punctate,
abdomen minutely so, sides of segments lightly pubescent. Post tarsi shorter
than usual, the basal segment scarcely as long as the rest combined. Dimensions:
10 x 4:5 mm,

Habitat—Mount Wilson, Blue Mountains (the author).

I took two examples, both, I think, 9, in January, 1929, that are quite distinct
from the two following species by colour and sculpture, though the three species
are allied. Holotype in Coll. Carter.

178 AUSTRALIAN COLEOPTERA. NOTES AND NEW SPECIES, Vili,

. Homotrysis SEXUALIS, N. sp.

Elongate-obovate; nitid black, lightly pubescent; antennae with basal segments
piceous, the apical reddish; tibiae red, palpi and tarsi pale red.

Head sparsely punctate, interspace of eyes less than the diameter of an eye
in g; antennae with segment 3 cylindric, 4-10 subequal in length, and consider-
ably widened at apex, 9-10-11 successively narrower. Prothorax rather convex,
apex arcuate, advanced in middie, base rather strongly bisinuate; widest at base,
thence lightly narrowed, anterior angles depressed and widely rounded off, the
posterior acute; lateral border unseen from above; disc showing medial depres-
sion towards base, and two large triangular depressions accentuating basal sinua-
tion; surface with large, round, rather close, punctures; these finer towards base,
a few short hairs perceptible at sides. Elytra slightly wider than prothorax at
shoulders, and more than four times as long; a narrow horizontal lateral border
seen from above on basal half; striate-punctate, the striae well-defined, and
containing punctures, large near base but gradually diminishing in size to apex;
intervals flat for the greater part, feebly convex on declivity; the striae bordered
by a single row of punctures of the same size as those in striae; the intervals
otherwise with sparse smaller punctures bearing short red hairs, especially at
sides and apex. Prosternum closely, meso- and metasternum very sparsely and
irregularly, punctate; the medial area nearly laevigate; abdomen finely and
sparsely punctate, with short recumbent hair. Dimensions: 12 x 4-5 mm.

Habitat—New South Wales: Oberon (Mr. F. H. Taylor).

I am indebted to my friend, Mr. Taylor, of the School of Tropical Medicine,
for this amongst other discoveries. A pair (J, 2) show striking sexual distinction,
but the differences are chiefly limited to the following characters: 9. The eyes more
wide apart, antennal segments less wide, the prothorax clearly wider towards
apex, giving a more robust, subquadrate appearance. The species is not very
near any other, being more elongate-obovate, the anterior sides of pronotum much
more rounded off than with H. laticollis Boh. Holotype ¢ and allotype 9? in Coll.
Carter.

HoOMOTRYSIS SILVESTRIS, N. Sp.

Elongate-ovate; sparsely pubescent, black, nitid; oral organs, antennae, tibiae
and tarsi red.

g. Head lightly punctate, eyes prominent and rather close, interspace less
than half the transverse diameter of an eye; antennae having 3-11 subequal in
length, 3 linear, 4-10 enlarged at apex, 11 very slender and pointed. Prothorax:
Apex truncate, base bisinuate, widest at base, thence lightly and arcuately narrowed
to apex, front angles depressed and rounded, hind angles acute; disc closely, not
densely, punctate with wide and rather deep triangular basal excisions, a few
upright red hairs at sides. Elytra as wide as prothorax at base, and more than
thrice as long; striate-punctate, intervals nearly flat, each with two rows of round,
clearly-cut punctures along the margins of striae; near base a few confusedly
scattered punctures, sides and apical region with recumbent red hair. Metasternum
strongly but sparsely punctate; abdomen finely so. Post tarsi with basal segment
longer than the rest combined. Dimensions: 9-10 x 3:5-4-2 mm.

Habitat——Queensland National Park, MacPherson Range (the author).

Three examples (2 g, 1 2) were taken by beating foliage in January, 1928,
and are notable for the unusually clear-cut sculpture, without rugae or wrinkles,

BY H. J. CARTER. 179

Narrower than H, sexualis (supra), the sculpture of upper surface is stronger, the
eyes closer, the antennal segments longer.

?. Eyes more distant, prothorax more widely rounded. Holotype and allo-
type in Coll. Carter.

HoMOTRYSIS MONTIUM, 0. SD.

This species is so near the preceding (silvestris) that at first it seemed
conspecific; but the following differences distinguish it: Colour subnitid brown,
legs (including femora) red, antennae and tarsi paler red.

dg. Eyes more widely separated, interspace about half the transverse diameter
of one eye; antennal segments short, stout and linear. Pronotum more finely
and closely punctate, the base only feebly bisinuate, with much less marked basal
excisions. Elytral intervals strongly but irregularly punctate. Episterna coarsely
punctate, abdomen longitudinally strigose. Post tarsi with basal segment about
as long as the rest combined. Dimensions: 3, 10 « 3; 9, 10 x 3-5 mm.

Habitat—New South Wales: Blue Mountains (the author).

A pair has long been in my cabinet unnamed.

©. More robust, the eyes wider apart. Holotype and allotype in Coll. Carter.

HYBRENIA BORCHMANNI, Nl. Sp.

Wide, obovate; nitid black, finely pilose; palpi, antennae and tarsi piceous,
legs dark.

Head and pronotum densely and finely punctate; clypeus with a red border,
labrum with a yellow fringe of hair; eyes large, prominent and rather close—
interspace less than half diameter of an eye; antennae narrowly linear, 3-11
successively shorter. Prothorax one and a half times as wide as long; apex and
base bisinuate, the latter feebly, the former clearly advanced at angles and at
middle; widest at base, sides lightly, arcuately narrowed to apex, anterior angles
depressed and rounded off, posterior obtuse; disc without obvious depressions,
punctures fine and close at middle, larger towards sides, the narrow lateral border
unseen from above, lateral half of disc with short, red, recumbent pile. Elytra
slightly wider than prothorax at shoulders, widest at apical declivity, narrow
herizontal border seen only near middle; striate-punctate, the striae lineate, with
small, closely-set punctures; intervals flat, wide and closely set with punctures,
in general larger than those in striae, round and separated on basal third, with a
tendency to transverse wrinkling and coalescence behind this; finely pubescent
at sides and apex. Underside glabrous and densely punctate, prosternum and
epipleurae coarsely so. Dimensions: 16 x 6 mm.

Habitat—N. Queensland: Endeavour River (in Coll. Borchmann).

A-specimen (4) was sent amongst other Cistelidae by my friend and fellow
worker, F. Borchmann, that is distinct from previously described species. In my
table (These Proc., 1929, p. 77) it could only be confused with H. pimelioides
Hope, a larger species with pronotum and elytra more coarsely and sparsely
punctate. Holotype in Coll. Carter.

Dimorphochilus.—Herr Borchmann has very courteously sent me for examina-
tion the actual female specimen of D. diversicollis Borch., photographed for his
paper (Faun. Sud. West. Aust., 1908). This confirms my opinion as expressed in
a former number of these PrRocEEDINGS (1932, p. 112) of its identity with D. gouldi
Hope. I would here express my high appreciation of this generous service to
the cause of accurate systematics.

186 AUSTRALIAN COLEOPTERA. NOTES AND NEW SPECIES, Viii.

Family CERAMBYCIDAER.
HESTHESIS DIVERGENS, N. Sp.

9. Narrowly elongate. Red with dark markings. Head black, thickly clad
with orange hair on front; antennae red, becoming obscurely dark from the 7th
segment; prothorax orange-red with a transversely oval, nude, black spot on
middle of disc; scutellum black, elytra orange-red, the external half (except
shoulders), also the apices reddish-brown; metasternum with yellow spot at
middle, sides and base; abdomen yellow—apical segment orange-red—with three
dark ventral and three dark dorsal bands. Prothorax roundly widened at middle.
Elytra sharply divergent from in front of middle, terminating externally in an
acute tooth, with small preceding incurvature. Dimensions: 19 x 4 (vix) mm.

Habitat—New South Wales: Mullaley (the author).

I took a single female example near Garrawilla homestead in November, 1932,
on Leptospermum flower. It has the elytra even more divergent than with the
western species, H. angulata Pase., from which it differs widely in colour. Its
orange apical segment of abdomen distinguishes it from all others of its congeners
except crabroides mihi and vesparia Pasc., from both of which it is readily
separated by its divergent elytra. The antennae extend to half the length of
the abdomen. ¢ latet. Holotype in Coll. Carter.

MICROTRAGUS DISCOSPINOSUS, nh. sp. Text-fig. 4.

Chocolate-brown, squamose, with long erect pale hairs on upper surface.
Elytra with two transverse white markings near middle; the apical region mottled
with pale areas.

Head impunctate, interspace between eyes in 9 less than the diameter of an
eye, in ¢ still narrower. Antennae strongly pilose, each segment with pale ring
at base; in the ¢ extending to apex of elytra, in 9 not quite so. Prothorax widest
at middle, sides lightly and evenly rounded; with two prominent, conical lateral
spines and two long recurved spines on disc; in two examples a minute tubercle
at middle, between the spines; surface pitted with large foveate punctures.
Seatellum small, triangular. Elytra ovoid, apices truncate, with strong recurved
basal spines, and five variously irregular rows of tubercles, placed more or less
en echelon, in those examples showing most regularity, the first two rows meeting
at base; surface pitted with large punctures, save where hidden by velvety derm.
Epipleurae with rows of smaller tubercles. Prosternum coarsely punctate.
Dimensions: § 10 x 32 mm.,?913x5 mm ©

Habitat—North Queensland: Endeavour River. National Museum and Coll.
Carter.

Four examples, 3 4, 1 9, before me are strikingly distinct by the presence
of the recurved spines on the disc of the pronotum instead of the usual tubercles,
and the truncate apices of the elytra. Holotype ¢ and allotype ? in the National
Museum, Melbourne.

CORYNEBACTERIA AS AN IMPORTANT GROUP OF SOIL MICROORGANISMS.
By H. L. JENSEN, Macleay Bacteriologist to the Society,
[Read 31st May, 1933.]

The genus Corynebacterium Lehmann and Neumann has been studied diligently
by medical bacteriologists, who naturally have devoted most of their energy to
the study of the numerous pathologically important organisms of this genus,
and very little attention has been paid to its purely saprophytic members.
Harris and Wade (1915) seem to have been the first to conduct a search
for corynebacteria occurring outside the human and animal body. They
found ‘“diphtheroids” of frequent occurrence, not only in various minor skin
lesions, but also in the air, and concluded as a result of their study that
“diphtheroids constitute a broader field of saprophytism than is generally appre-
ciated’. Kisskalt and Berend (1918) considered that several non-spore-forming
bacteria, e.g., Bact. helvolum and Bact. erythrogenes, should really be included
in Corynebacterium because of their characteristic mode of cell division. This
classification has been adopted in the most recent edition of the manual of
Lehmann and Neumann (1927), who also isolated such organisms from air and
water.

The main points in the definition of Corynebacterium are: non-motile bacteria
without endospore formation, gram-positive, not acid-fast, generally rod-shaped,
but with a marked tendency to formation of irregular, club- or wedge-shaped,
sometimes branching cells of a more varying size and shape than is usually
found among the Hubacteriales, and multiplying by a characteristic “snapping”
division of the cells, which causes the bacteria in microscopical preparations to
appear in V- or III-like arrangements, or irregular groups sometimes compared
to Chinese letters.

In a number of microbiological analyses of Australian soils the writer became
aware of a remarkably frequent occurrence of bacterial colonies of this type on
agar plates. An estimate of their relative frequency was carried out in the
following manner: platings were made from soils in adequate dilutions
(1:100,000 —1,000,000) on 4-5 parallel plates of dextrose-casein-agar*; after incuba-
tion for 8 days at 28-30° C., all bacterial colonies were counted, and 40-50 colonies
were picked at random and examined microscopically in nigrosin smears, by
which method of preparation the characteristic cell shape and angular arrange-
ment of the corynebacteria appears very clearly. The percentage of
corynebacterium-like colonies was then calculated. Colonies of “doubtful”
character, as were sometimes met with, were not reckoned as corynebacteria.
From each soil a number of strains (generally 3-4) were isolated from typical
colonies and studied in pure culture; they all conformed with the above definition
of Corynebacterium. A closer description of the morphology and biology of these
organisms is reserved for a later occasion.

* Dextrose 2:0 gm.; casein, dissolved in 10 c.c. 0-1n NaOH, 0:2 gm.; K,HPO, 0°5 gm.;
MgSO, 0-2 gm.; agar 15-0 gm.; water, 1,000 c.c.; pH 6:5-6-6.
E

182 CORYNEBACTERIA AS AN IMPORTANT GROUP OF SOIL ORGANISMS,

TABLE 1.

Occurrence of Corynebacteria in Various Soils.

Corynebacteria.
pH Bacteria.*
Soil Character of Soil. of Millions
No. Soil. per gm. % of total Millions
Bacteria. per gm.
la Heavy loam, rich in organic matter, grass
land, Sydney University. 4°8 14-4 29 4-2
1b Same with addition of 1% cellulose and
0:1% ammonium sulphate, kept moist
5 months, room tpt. 4-2 0:6 8 0-05
le Same as la, with addition of 2% calcium
carbonate, kept moist 14 days. 7:4 99-0 29 28-7
2a Very light sand, poor in organic matter,
vacant grass land, Bellevue Hill, Sydney. eal 8-0 37 3:0
26 Same with addition of 1% calcium carbonate,
kept moist 15 days, room tpt. Gor 10:0 48 4-8
3 Coarse sand, poor in organic matter, Bathurst,
N.S.W. 5-1 6-1 26 1-6
4 Light sand, rich in organic matter, Cooper
Park, Sydney. 5:3 17-3 13 2°3
5 Heavy loam, rich in organic matter, from
flower bed, Sydney University. 5:6 26-3 35 9-2
6 Light loam, very rich in organic matter,
from Scone, N.S.W. 6-0 17:9 39 7-1
7 Light sand, poor in organic matter, Bellevue
Hill, Sydney. 6-1 10-0 33 Bon}
8 Red loam, poor in organic matter, Griffith,
N.S.W. 6-1 18-2 64 11-6
9 Heavy loam, rich in organic matter, from
pot experiments, Sydney University. 6:6 25-0 39 9-8
10 Loam, rich in organic matter, North Sydney. 6-9 20-2 60 i751
11 Clay, rich in organic matter, sheep pen,
McMaster Laboratory, Sydney University. este, 22-4 45 10-1
12 Loam, rich in organic matter, flower bed,
Sydney University. 7:3 25:3 65 16-4

The figures in Table 1 show plainly that the corynebacteria account for a
quite considerable part of the flora appearing on the plates, in some instances
(soils 8 and 12) even up to two-thirds of the total number of bacterial colonies.
There is a certain indication, although not very marked, that the percentage of
corynebacteria increases somewhat with decreasing soil acidity. In the very
acid soil, No. 1b, their frequency is remarkably low; this agrees with the fact
that a degree of acidity corresponding to pH 4:3-4:6 appeared critical for most
strains in pure culture.

Comparatively little work has yet been carried out on those non-spore-forming
soil bacteria which do not distinguish themselves by striking physiological
functions, such as nitrification, nitrogen fixation, denitrification, cellulose decom-
position, etc. One of the few bacteriologists who have made a real study of the
“ordinary” soil bacteria, viz., Conn (1925, 1928), describes the bulk of these as
“slow-growers” or “punctiform-colony-forming bacteria’, characterized by a poor

*Not including actinomycetes.

BY H. L. JENSEN. 183

growth and uncharacteristic fermentation reactions in liquid media. Conn’s
groups III and IV seem to have something in common with the corynebacteria,
but they have not yet been described in detail and were stated to be compara-
tively rare. The “slow-grower” which Conn found of most common occurrence—
Bact. globiformis—was not met with in the present investigation, and upon the
whole the corynebacteria dealt with here show little resemblance to Conn’s “slow-
growers”, since they all grew well in broth as well as on agar media, and gave
characteristic fermentative reactions. Most of their colonies on dextrose-casein-
agar were of a fair size, from 0:5 to 2 mm. in diameter after 7-8 days, white to
yellow, opaque, with a well-defined, even outline, of a mostly soft-pasty, some-
times slimy consistence; surface colonies were usually round, smooth and convex,
deep colonies lens-shaped or ellipsoid. Probably we must here seek the identity
of those soil bacteria which Greig-Smith (1911) interpreted as Rhizobia—nodule
bacteria of leguminous plants. The work of later investigators has not revealed
such large numbers of rhizobia in soil, and the nitrogen fixation recorded by
Greig-Smith cannot be regarded as proving the organisms to be rhizobia.* On
the other hand, Greig-Smith’s description of the appearance of the colonies and
the organisms in them agrees remarkably well with the corynebacteria in question.
It is well known that nodule bacteria at a certain stage of their life cycle produce
irregular, swollen and partly branched forms which may resemble corynebacteria,
and this suggests that Greig-Smith may have mistaken such organisms for
rhizobia. In order to test this point, platings were carried out from a few of
the soils in Table 1 on the levulose-asparagine-citrate-agar employed by Greig-
Smith, and the frequency of colonies of corynebacteria was estimated as before.
Numerous corynebacteria were found to develop on this medium, as shown below:

Percentage of corynebact. :
Bacterial colonies,
Soil No. Dilution. average per plate.
Levulose agar. Casein agar.
2b 1: 25,000 11-8 15 48
9 1: 50,000 18-3 24 39
12 1: 100,000 22-0 50 65

As a further test, 9 strains of corynebacteria known to be able to utilize
asparagine were grown on slopes of Greig-Smith’s agar, where they all, after 3
Gays at 28° C., produced a fair to abundant growth, convex, smooth, glistening,
opaque and whitish, not unlike that of nodule bacteria on some favourable agar
medium. Microscopical examination showed the presence of all the cell types
which Greig-Smith describes thus: “... irregular outline and structure suggesting
a sausage-skin stuffed more or less with marbles; and although the y and Y forms
were rare, the exclamation mark (!), the irregularly divided rod, and the club-

* Critical studies by several recent investigators have shown conclusively that
nodule bacteria do not fix elementary nitrogen when grown outside the host plant, but
several sources of error may give rise to a spurious nitrogen fixation.

184 CORYNEBACTERIA AS AN IMPORTANT GROUP OF SOIL ORGANISMS,

shaped form were quite numerous.”

Smith’s “rhizobia” were really corynebacteria.
be rhizobia is out of question; none of them shows anything like the characteristic
life cycle which distinguishes the nodule bacteria (Bewley and Hutchinson, 1920).

Some experiments were carried out in order to get an idea of the possible
importance of the corynebacteria in the decomposition processes in the soil.
200-gm. portions of the soils 2b and 9 in Table 1, with 10 and 25 per cent. of
water, respectively, received the following additions of organic matter:

0:-5% dried and ground mycelium of Penicillium sp. (“glaucum’’).

0:5% dried and ground leaves and stems of white clover.
1:0% dried and ground oats straw.

TABLE 2.

Multiplication of Corynebacteria during Decomposition of Organic Matter.

Soil and Addition.

Sand soil 2b, with 0:5% mycelium of Peni-
cillium sp.

Sand soil 2b, with 0:-5% dry matter of white
clover.

Sand soil 26, with 1:0% oats straw.

Loam soil 9, with 0:5% mycelium of Peni-
cillium sp.

Loam soil 9, with 0°5% dry matter of white
clover.

Loam soil 9, with 1:0% oats straw.

All this renders it very likely that Greig-
That the present organisms should

Corynebacteria.
Bacteria, Actinomy-

Time. Millions cetes,

per gm. % of Millions Millions

Bacteria. per gm. per gm.
Start 10 48 4°8 32
4 d. 1,333 13 173°3 (0)
8 d. 550 13 Cakets) 0-5
14d PABM | 38 89-1 alos)
22 1d 63 35 22-1 1-2
Start 10 48 4°8 3:2
4d. 765 27 206°5 0:3
8 d. 469 19 89-2 0-8
14 d. 170 23 39-1 1-2
22 d. 59 26 15-3 1°8
Start 10 48 4:8 3°2
4d. 95 35 33°3 0:9
8 d. 86 32 27°5 11983
14d 97 oo 32:1 1-2
22d 41 30 12:3 0°8
Start 31 38 11-6 aloat
4d. 352 12 42-2 4-0
8 d. 328 13 42-7 4-3
15s 212 27 57°3 4-5
22 d. 135 28 37-9 5-8
Start 31 38 11-6 Thoal
4d 264 25 66-0 2-0
8d 272 25 68-0 3:8
15d 194 28 54-4 5-3
22 d 139 20 27-8 3:7
Start 31 38 11-6 1-1
4d. 494 17 84-0 2°8
8 d. 345 29 100-0 4-3
15d 235 35 82-4 4-8
22d 210 32 67-1 2:4

BY H. L. JENSEN. 185

The soils were kept in large Petri dishes for 22 days at room temperature
(18-20° C.). During this period, plate counts of bacteria were carried out four
times, and the frequency of corynebacteria was estimated. The results, reproduced
in Table 2, show that not only do the corynebacteria occur as a constituent of
the soil flora, but they also multiply strongly during the decomposition of organic
matter and thus presumably take a part in these processes. Their percentage is
generally highest during the later stages of decomposition, and higher in soils
with straw and clover than in soils with fungous mycelium, where a motile,
yellow bacterium, probably Bact. herbicola or a related form, flourished abundantly,
especially after 4-8 days.

SUMMARY.

Bacteria possessing the characters of the genus Corynebacterium were found
to occur as a numerically important group of microorganisms in Australian soils,
accounting for 8 to 65 per cent. of the numbers of bacterial colonies developing
on plates of dextrose-casein-agar. They appear to be active in the decomposition
of organic matter in soil, particularly in the later stages of the process. They are
probably identical with certain organisms previously recorded as rhizobia.

References.

BEWLEY, W. F., and HUTCHINSON, H. B., 1920.—On the Changes through which the
Nodule Organism (Ps. radicicola) passes under Cultural Conditions. Journ. Agr.
Sci., 10, 142-161.

Conn, H. J., 1925.—Soil Flora Studies. VI. N.Y. State Agr. Hxup. Stat. Techn. Buil.,
INO wuli5s

————,, 1928.—Certain Abundant Non-Spore-Forming Bacteria in Soil. Cent. f. Bakt.,
II, 76, 65-88.

GREIG-SMITH, R., 1911.—Contributions to Our Knowledge of Soil Fertility. II. Proc.
LINN. Soc. N.S.W., 36, 492-503.

Harris, W. H., and Wapbsr, H. W., 1915.—The wide-spread Occurrence of Diphtheroids
and their Occurrence in Various Lesions of Human Tissues. Journ. Exp. Med., 21,
493-509.

KISSKALT, K., and BEREND, E., 1918.—Untersuchungen itiber die Gruppe der Diphtheroiden
(Corynebakterien). Cent. f. Bakt., I. Orig., 81, 444-447.

LEHMANN, K. B., and NEUMANN, R. O., 1927.—Bakteriologische Diagnostik (7th Ed.,
Munich. English Translation, New York, 1931).

THE MARINE PLANKTON OF THE COASTAL WATERS OF NEW SOUTH
WALES. I.
THE CHIEF PLANKTONIC FoRMS AND THEIR SEASONAL DISTRIBUTION.

By Proressor WILLIAM J. DAkin, D.Sce., F.Z.S., and ALLEN CoLerax, B.Sc.,
Zoology Department, University of Sydney.

(Plate vii; seven Text-figures.)
[Read 28th June, 1933.]

Introduction.*

Since this paper is the first of a series of plankton studies which it is intended
should come from the Zoology Department of the University of Sydney, and the
associated Sydney University Biological Station, it is desirable that the general
aim of the work should be explained at the outset.

Systematized studies of the biology of the ocean waters off the coasts of
Australia can only be said to have been initiated very recently. It is true that
there are some apparent exceptions to this statement. Various expeditions have,
from time to time, collected in Australian waters, dating from the ‘Challenger’
Expedition in 1874 to the ‘Dana’ Expedition in 1929. Local expeditions, more
especially from Sydney, have brought in vast quantities of dredged materials, and
thanks in particular to the activities of the Australian Museum, the systematic
zoology of the Australian coastal areas is very well known. A rather elaborately
organized expedition spent a year on the Barrier Reef in 1928-1929, and carried
out a very thorough biological investigation of the chosen base.

Notwithstanding this, however, it is surprising that until recently there
was no marine biological station in the whole of Australia—even at present there
is only one small semi-private station in occupation—and no organized work
at sea on the plankton or the marine hydrography has been carried out, apart
from that of the Barrier Reef Expedition. Again, whilst a systematic study
of the Australian fishes has been pursued very satisfactorily indeed, practically
no serious researches into the general biology of the fish fauna have ever been
made.

To return to the matter of the planktonic life, our Knowledge even of the
systematics is confined to published accounts of a very few catches taken in
Australian waters, by expeditions which were en route to other regions—the
“Challenger”? made a few catches in the coastal waters, and the British Antarctic
Expedition (‘Terra Nova’), 1910-1913, made tow-nettings whilst between New
Zealand and Australia in the winter of 1911. The German surgeon, Kramer,
used a tow-net for Copepoda whilst in New Zealand coastal waters, at Port
Jackson, and again at Melbourne.

Tow-nettings have, of course, also been frequently made for teaching purposes
—by members of the staffs of different Australian Universities, and by others
interested, without any serious attempt being made to diagnose the species
obtained.

* The pursuit of these marine investigations has been valuably aided by grants
from the Research Endowment Fund of the Council for Scientific and Industrial Research
and also the National Research Council of Australia, to whom our thanks are warmly
tendered.

BY W. J. DAKIN AND A. COLERAX. 187

It was only the provision by the University of facilities for the continuous
collection of plankton and water samples off the coast near Sydney which enabled
us to initiate a long period investigation, the chief aims being to discover (1) the
seasonal variations in the plankton both qualitatively and quantitatively, (2) the
relation of plankton changes to the physical environment, and (3) the eggs and
larvae of the food fishes of New South Wales waters and the conditions under
which they are spawned and hatched. We are only beginning now to turn our
attention seriously to the third of these aspects, for the initial difficulties in
opening up a comparatively new field necessitated a considerable expenditure of
time on taxonomic plankton studies until we became familiar with at least the
organisms most commonly present.

MrErHoDs AND STATIONS.

It seemed better at the outset, that the investigation should be concentrated
at one station and under conditions which would be as little complicated by
estuarine waters as possible. With this in view, a point was selected four miles
east of North Head, Port Jackson, Sydney, and efforts were made to reach this
spot at fortnightly intervals, at or about the hour of high water, and at about the
same time of day on each occasion. The choice of high water was simply to
ensure that for five or six hours ocean water had been flooding into Sydney
- Harbour, and we could count definitely on no contamination from that source.
It is probable, however, that even at the hour of low water on most occasions we
should have been out of the range of ebbing harbour water, which usually moves
in a southerly direction rather than northerly, once it has escaped from the
harbour.

Although this procedure has been the basis of the main scheme, collections
have been made at odd times at other localities, and also on special occasions
from trawlers many miles to the south of our station. A few collections have
been made at night, with a view to comparing day and night hauls at the same
station.

Complete records have been made of the weather and, before taking the
plankton catches, a series of water samples were often collected, together with
temperature readings at various depths from the surface to the bottom. When
this was not done, the station was again visited for this purpose within 24 hours.
It might be regarded as better to make hydrographic observations at the same
time as the plankton catches, but, in this connection, it must be observed that
we were working in unsheltered ocean waters from a small craft and usually
under considerable difficulties. As it was, we had to experiment and construct
gear to simplify the work with the water bottle in consequence of the rolling and
pitching which was almost constantly our lot. Moreover, our early work showed
little change at our Station, whether in the planktonic or the hydrographic
observations during 24 hours, and it must be remembered that at many places
elsewhere physico-chemical observations for the study of seasonal variations
during the twelvemonth are only made at monthly intervals. For a considerable
period, we conducted hydrographic observations at sea once a week.

The plankton samples have been taken with nets of both silk and cheese-
cloth with a mesh rather like Stramin.* For the continuous work, we used
conical nets 32 inches in length, with a diameter at the mouth of 14 inches. The
silk part of the cone was 26 inches in length. Two nets, fine and coarse, were

* Recently we have used imported Stramin.

188 MARINE PLANKTON OF THE COASTAL WATERS OF N.

always towed horizontally at the surface for ten minutes,
whilst another coarse net was towed horizontally at a
depth of approximately 30 metres for the same time. In
addition, catches were made, both at the surface and at
different depths, with a large cheese-cloth net (mouth 30”
in diameter), but these catches were not used quanti-
tatively.

Qualitative catches were also made by vertical hauls
with a large net of the same type and size as one of the
“Discovery 2nd” (Whaling Exploration) nets, but these
had to be discontinued owing to the difficulty of using
this large size net. In addition to the above, two cylindrical
nets were made of bolting silk, approximately 25 feet in
length, but only 6 inches in diameter. They were long
slender cylinders, and were designed so that they could be
hauled at full speed (7-8 knots). They worked excellently,
but probably our speed rarely exceeded 6 knots when we
were out. These nets were towed over a long distance and
the contents examined solely for qualitative studies of the
plankton.

It is a great pity that no standard nets of reasonably
small size have been designed and agreed upon inter-
nationally. There is, it is true, a Standard Helgoland
Net, but this was designed for vertical hauls, and in any
case is not, so far as we are aware, in general use. It is,
however, fairly obvious that, for accurate quantitative
comparisons of the plankton production of different places,
more accurate standards than those of hauls with silk
nets are absolutely essential. Another point which is not
sufficiently stressed in this connection is the need for
using a much greater diversity of nets, and of size of
mesh, if a true picture of the plankton is to be obtained.

It was soon realized that only one method could
reasonably be adopted for estimating quantitatively the
components of our catches—that in which the different
organisms are counted in a fraction of, or in the whole
catch. In other words the Hensen methods were applied,
and although tedious and tiring, they proved as usual
that mere estimation that a species is common, very
common, or rare, is of little or no value in the accurate
determinations of seasonal variation. Counts were made
in samples of 0-1 c.c. and 0-5 c.c. of fluid taken from such
dilutions of the catch as proved suitable. The larger
organisms were counted in the undiluted catch. The small
quantities of fluid were extracted by special pipettes after
complete shaking of the whole catch, and the sample was
estimated on a ruled plate under the microscope.

THE HyproGRAPHIC CONDITIONS.
It has been deemed advisable to describe and discuss
‘the physico-chemical conditions which prevail in the ocean

S. WALES, i,

a
41 Marsch

JAN. G21 Feb.

Y
wo RR) Gp tS be

Text-fig. 1—Sea tem-

peratures at the station

4 miles east of North

Head, during 1931 and
part of 1932.

BY W. J. DAKIN AND A. COLEFAX. 189

waters from which the plankton has been taken, in another paper which will
appear shortly after this. Reference to some of the data will, however, be made
in our general discussion. It has been thought advisable to reproduce the chart
of the sea temperatures here since it is of prime importance in visualizing the
range of seasonal change in our locality. A brief account of the chief forms
present in the plankton will first be given and then the seasonal distribution of
the more important types will be described and discussed.
Tue Driaroms.

The study of the diatoms was forced upon us by the prime necessity of deter-
mining the seasonal changes amongst the “producers” of the plankton. It is not
the first time by any means that marine zoologists have been compelled to take
the marine diatoms into their purview. From the nature of things we have,
therefore, treated the diatoms in a more general manner than the zooplankton.

The diatoms were chiefly caught in the finest meshed nets and it was in the
quantitative hauls with such nets that we looked for seasonal changes in plankton
production of the kind formerly so familiar to one of us (W.J.D.) in the catches
round Great Britain. Only on one or two occasions, however, could one speak of
really big catches, and probably on no occasion did a net come in coated with
the thick brownish or green slime so characteristic of hauls during the diatom
maximum in the seas mentioned.

So much work has been carried out on the marine diatoms of the Northern
Hemisphere that it was deemed advisable to send samples from our catches for
direct comparison with the northern forms. Fortunately very able studies on
the diatoms have been made by Professor W. E. Allen, of the Scripps Institute
for Oceanography, California, and it was only natural, therefore, to send to that
Northern Pacific Station for assistance. We are happy to acknowledge the help
unstintingly given by Professor Allen.

The following list represents only a few of the species present, but it includes
those which play the most conspicuous part in the make-up of our plankton as
a whole. In certain cases where counting of individual species of a genus would
have involved much labour and difficulty, the species are lumped together under
the head of the genus.*

Family CoscINopIsScAcEAE Schroeder.

Genus Coscinodiscus Ehr.—The only species of this genus, indeed of the
family, which we have separately recorded, is Coscinodiscus concinnus, a large
diatom of wide distribution recorded from North Atlantic waters and from the
Northern Pacific off California. Its abundant occurrence is practically confined
to the spring months (see tables).

Coscinodiscus Group.—Under this head we have grouped together in the
tables not only species of Coscinodiscus, but of other genera of the family
Coscinodiscaceae, viz. Asteromphalus. Further specific studies involving com-
parisons with types will be necessary before a complete list of the species occur-
ring can be given. With the exception of the genus Planktoniella, the group has
not played a great part in the make-up of the phytoplankton during the period so
far studied.

* Since this paper was written, we have received a further communication from
Professor Allen and Miss Cupp embodying a list of some of the species which are more
rare than those entering into our quantitative estimations (some of them have been
included in our counts but not recorded separately). A more complete systematic list
of the diatoms will be given in a subsequent paper.

F

190 MARINE PLANKTON OF THE COASTAL WATERS OF N. S. WALES, i,

Planktoniella sol (Wallich).—This is an oceanic plankton species of unmis-
takable appearance. It occurs in the North Sea and the North Atlantic, but is
noteworthy in tropical waters. It was most abundant at an unusual time for the
diatoms—the months of June and July, 1931, and in July, 1932, one of the days.
gave almost nothing but Planktoniella in the fine net catches.

Family THALASSIOSIRACEAE Lebour.

Genus Thalassiosira Cleve—Four species of Thalassiosira have played a
prominent part in the make up of the phytoplankton, and three of these, 7. rotula,
T. gravida, and T. condensata, evidently require the same conditions, for they
tend to occur in abundance at the same time. The spring maximum is sharply
defined and commences in the middle of August or thereabouts, extending into
October.

Thalassiosira rotula Meunier.—This species, recorded as a temperate form, is
known from the Atlantic off France, and from the Pacific off California.

Thalassiosira gravida Cl.—This species, so well Known in the Northern Hemi-
sphere (it is one of the common diatoms of British seas and is recorded from
the Northern Pacific), occurred together with 7. rotula, which exceeded it in
numbers. The two species were for convenience counted together.

Thalassiosira condensata (Cleve).—This diatom appeared in huge numbers.
in the same weeks of 1931 and 1932. It is easily mistaken for Lauderia, as several
writers have pointed out. Its distribution is also extensive—North Atlantic as
well as North Pacific (off California).

Thalassiosira subtilis (Ostenfeld).—Much less common than the other forms.
Recorded during June, July and August.

Family SKELETONEMACEAE Lebour.

Stephanopyzis sp.—During a very limited period in the springtime of each of
the years 1930-1932, an enormous influx of the genus Stephanopyzis occurred. Two
species were found together in this “‘explosive’’ maximum, the most common one
being S. turris, the other S. Palmeriana. Stephanopyzis, so far as we know,
does not seem to have been recorded anywhere else as one of the dozen or less.
species which make up the bulk of the plankton at the times of maxima. NS. turris
is usually regarded as an Atlantic species. S. Palmeriana has a wider range.
Their massive occurrence in our waters is particularly interesting since no less.
than four species were recorded in a catch made by the ‘Challenger’ in the
Arafura Sea (North of Australia).

Curiously enough, the related genus Skeletonema, which is one of the most
abundant diatoms of the spring maxima of British seas and is regarded as one
of the leading half dozen of the Californian Coast, is relatively rare with us.

Skeletonema costatum (Grev.) Cl.—This species, as noted above, is charac-
teristic of Northern Seas; it occurs spasmodically in our catches and was present
between May and September.

Family LeprocyLINDRACEAE Lebour.

Dactyliosolen mediterraneus Peragallo (D. tenuis (Cleve) ).—This North
Atlantic, Indian Ocean and North Pacific form is never very common, but occurs
at odd times almost throughout the year.

Leptocylindrus danicus Cleve.—This diatom was most common during 1931-32
in the summer time. It frequently occurs in large numbers. It has been recorded
from Arctic Seas, North Atlantic, U.S.A. Coast of Pacific, Malay Archipelago and
Japanese waters, also from Antarctic Seas.

BY W. J. DAKIN AND A. COLEFAX. 191

Leptocylindrus minimus Gran.—This species is also common at times,
generally occurring with L. danicus.

Guinardia flaccida Castr.—This species, of wide range, has occurred at odd
times, but has not taken part in making up obvious diatom maxima. It is
recorded in a surface catch made by the “Challenger”, in the Arafura Sea.

Family CorerHroNACcEAE Lebour.
Corethron eriophilum Castr.—This species, recorded from the Arctic, North
Atlantic and North Pacific, has been frequently noticed in the plankton. Quanti-
tatively it is insignificant.

BACTERIASTRACEAE Lebour.
Bacteriastrum sp.—Species of this genus are occasionally present, but quan-
litatively the part played by this genus is insignificant.

Family RHIZOSOLENIACEAE Schroeder.

At least ten or more species of Rhizosolenia have been noticed in our catches.
Those which play a really big part in the constitution of the diatom plankton are
the following:

Rhizosolenia alata Brightw.—This is probably our most common species of
Rhizosolenia, It is an oceanic diatom of wide range and common occurrence in
the world’s seas. It extends from sub-Arctic waters to the Antarctic, is found
in the Atlantic, Pacific and Indian Oceans, and has even been recorded from the
Black Sea.

R. stolterfothii H. Perag.—This species is often very common in our plankton
catches and must be ranked as quantitatively important. It does not, however,
form such definite maxima as other Rhizosolenia species. It ranks as one of the
most common species in British Seas.

R. robusta, R. setigera, R. calcaravis, R. styliformis and Rk. imbricata have all
been noted in the plankton at different times.

Seasonal distribution of the genus is referred to in the general discussion
which follows this section of the paper.

CHAETOCERACEAE Schroeder.

Chaetoceras sp.—The species of this genus may almost be said to play the
most important part in the matter of bulk in the phytoplankton. For convenience
in counting, all but one species have been lumped together in our quantitative
studies. It is scarcely possible to obtain any diatom catch without Chaetoceras
being present. The quantity increases, however, considerably during the spring-
time and, in 1931-32, during the late summer (see notes in general discussion).

The following species have been distinguished: C. afinis Laud., C. Lorenzianus
Green, C. Peruvianus Gran., OC. didymum Ehr., C. curvisetum Cleve, and C. debilis
Cleve. One of the most constant is C. curvisetum. :

C. sociales Lauder.—This characteristic form, in which the small chains are
held together in gelatinous colonies, has presented a sharply defined maximum in
the early spring (August 2nd-31st) of 1931. The species has been recorded from
Arctic Seas, from the North Atlantic, North Sea and from Japanese waters.

BIDDULPHIACEAE Lebour.
Biddulphia mobiliensis (Bail.) and Biddulphia regia Schultze.—There is one
common Biddulphia species in the New South Wales plankton and it was originally
named as B. mobiliensis, which is probably the most common species of the

192 MARINE PLANKTON OF THE COASTAL WATERS OF N. S. WALES, i,

plankton collections made elsewhere in the world. It is now apparent, however,
that B. mobiliensis and B. regia have frequently been confused. Our specimens
are most like B. mobiliensis of Gran. (Nordisches Plankton). If this is actually
B. regia, the form collected in our plankton must be designated under that
name. Allen, however, regards it as B. mobiliensis.

Biddulphia mobiliensis looms large in the diatom catches of the spring
months, August and September. No other species of Biddulphia has been
diagnosed with certainty and, in any case, mo other species is of consequence from
the quantitative aspect.

Hemiaulus Hauckii Griin—Occurs at odd times in small numbers but may at
times become very abundant, as in the catch of November ist, 1931. It is known
from the North Atlantic and North Pacific Oceans, as a neritic species.

Ditylliium Brightwelli (West.).—This diatom, whilst it may occur sporadically
in various months of the year, develops a distinct, and quantitatively an important
maximum in springtime (August and September).

It is another neritic planktonic species which is of wide distribution—coasts
of England and North Sea, North Atlantic, and Japanese Seas.

EUCAMPIACEAE Schroeder.

Eucampia zoodiacus Ehr.—This is an important constituent of our coastal
plankton. It occurs over a wide range of months, and presents a marked maximum
in the springtime. In 1931, the greatest abundance occurred as late as November.
In Californian waters, where it is very common, it also occurred at any time of
the year, and the same thing applies to its presence in Japanese waters. The
species is a neritic form found in the North Atlantic and North Pacific, but not
extending into Arctic Seas. It is possible that a second Hucampia species
occurring in much smaller numbers may have been counted in with the above
species, but further investigation will be necessary to determine this.

Climacodium Frauenfeldianum Griin.—This diatom, recorded by British
workers as an oceanic and tropical Atlantic form, is of frequent occurrence. It is
significant that the greatest abundance is recorded for the later summer months,
when the sea temperature is at its highest. It would appear, therefore, that in
the Southern Pacific, its correlation with high temperature water is equally well
demonstrated.

Streptotheca thamensis Shrubs.—In contrast to Climacodium, this “chain
forming” species is neritic and, whilst occurring sporadically throughout the year,
presents a maximum which commences in the later winter months just before
spring. Also in contrast to Climacodium, it is recorded in Hurope as an Arctic
species. It is distinctly interesting to see how these correlations with environment
recorded in one hemisphere are now borne out in another. A different species
of the genus has been recorded from the Indian Ocean.

FRAGILARIACEAE Schroeder.

Asterionella japonica Cleve and Moller.—This species is one of our most
abundant types. It occurs over a wide range of the year, but is particularly
abundant in the spring and presents a second and smalier maximum in the late
summer (see tables). In Japanese waters it attains its maximum in late autumn
and winter.

This diatom is a neritic form and of wide distribution, having been recorded
from the coasts of European countries, the North Atlantic, North Pacific off Japan,

BY W. J. DAKIN AND A. COLEIPAX. 193

off California where it is one of the most abundant species and from the Indian
Ocean. At times it occurs in swarms—this fact noted in European waters holds
good also in the New South Wales coastal region. ;
Thalassiothrix nitzchioides Grin.—The species occurred at odd times during
the year, but was most common in September.
Thalassiothrix longissima Cleve and Griin.—Of less importance than
T. nitzchioides. The only considerable catch appeared on 23rd April, 1932.

Family NirzscHrackag. .

Nitzschia seriata Cleve.—This is a common species in New South Wales
waters. It is of frequent occurrence in almost all months of the year and is
probably always present, even though rare at times. It presented a maximum in
the late summer months of 1931 and in the summer of 1932. It also presented
a smaller maximum in the spring. It has been recorded at its greatest abundance
in Japanese waters in the spring months. The distribution of this diatom is
absolutely cosmopolitan—it occurs from the Arctic Seas to the Antarctic. It
is termed neritic by Lebour, but its distribution does not seem to be confined
to neritic regions.

Nitzschia delicatissima Cleve = N. longissima Ralp.—This diatom has been
recorded, but is relatively unimportant quantitatively. It is a neritic form,
recorded from the North Pacific and from the Indian Ocean, as well as from the
European Coasts and North Atlantic.

Nitzschia closterium (Ehr.) Sm.—The species, though not abundant, has
been recorded sporadically in different months. It is unimportant quantitatively.

FurtHER Nores 0N THE PLANKTON DIATOMS.

It will be seen from the notes given above, that the diatom plankton of
New South Wales waters, so far as our records show, consists of the usual
admixture of oceanic and neritic species, which might be expected within the
vicinity of a continental shelf and the open ocean and, as usual, only a few
species are quantitatively of great significance. Neritic species are those which
show some sort of dependence on the presence of land or of shallower water.
The dependence may be due to the fact that resting spore stages must rest on
the sea bottom, or that other requirements are not met in the open ocean.

All the species which are likely to be of importance as “producing” organisms
(contrasted with the animal plankton—the “consumers’’) are well known types,
and of wide distribution in the seas of the Northern Hemisphere. The present
records, however, extend the known range of distribution for practically every
form quoted.

It is interesting to notice that, whilst peculiarly Antarctic species play
little part in our catches, there is an admixture of temperate and tropical types.
The invasion of Planktoniella sol, which sometimes occurs, is particularly
interesting in this respect. The diatom facies reflects the fact that we are
working on the edge of a tropical current and in a neritic zone. Comparisons
of our diatom plankton with that from the Tasmanian region would be most
interesting. It might be useful to remind readers of the fact that diatoms play
a most important part in neritic phytoplankton because of their power of
developing amazingly when the requisite food stuffs are richly present. Coastal
waters are, compared with the open ocean, richer in such food stuffs. The same
thing applies to certain areas of the sea in high latitudes.

194 MARINE PLANKTON OF THE COASTAL WATERS OF N. S. WALES, i,

Tue SEASONAL DISTRIBUTION OF THE DiATomMs IN New SoutH WALES WATERS.

The quantitative and qualitative changes in the plant life of the plankton is
of outstanding importance, for it is the fundamental indicator of productive
rhythm in the sea. Now the change in volume of the diatom catches during the
year at Sydney is nothing like so marked as it is in certain colder seas. S2asonal
variations were scarcely obvious from inspections of the nets. It was not until
our enumerations were complete that a picture of diatom change directly compar-
able with that of Northern Seas became clear. For purposes of comparison,
curves showing the seasonal distribution in the Irish Sea as well as in our
waters are given in Text-figures 2 and 3.

Jan feb. Mar. Apr May Jun “Jul Aug Sep Ocl Nov Dec.

“ \

Sei attesmes ens oe oN Ses ee ne
Fe a
ae net Apr. May June July Aus Seb. Oct Nov. Dee.
nie ~SihdHlagettate

Text-fig. 2.—Diatom production in New South Wales coastal plankton, 1931.
Text-fig. 3.—Curves indicating the seasonal abundance of the chief
constituents of the plankton of the Irish Sea (from a paper by Herdman).

During the winter months of May, June and July, the diatoms, whilst never
absent, were most reduced in number. Then, at some date in the month of
August, a change occurred, heralded by an increase in Chaetoceras species, in
Asterionella japonica and Thalassiosira. Thalassiosira condensata apveared
almost “explosively” in the two consecutive years under review, at approximately
the same date in August. After six weeks or so Asterionella began to diminish
again (at the end of September), and the Thalassiosira maximum was of about
the same duration, but species of Chaetoceras kept the diatom maximum still up
until the month of October was through. Following on the genera mentioned,
Rhizosolenia increased in numbers and attained its maximum in November, 1931,
after most other diatom species had commenced to wane. This marked the end
of the vernal maximum. During the last weeks of November and in December
the plankton was very poor in diatoms. January, however, brought a recrudes-
cence, mainly due to Leptocylindrus, Rhizosolenia, and Chaetoceras, and this

BY W. J. DAKIN AND A. COLEIAX. 195

continued into what might be called a small autumn or late summer maximum
with Asterionella japonica appearing again.

The net result is that a curve representing the rise and fall in the quantity
of producing plankton presents a peak in early spring and another smaller one
in late summer. ;

One of the most striking results of this, our first work, is to demonstrate a
similarity with the plankton rhythm of European Seas. The abundance of plank-
tonic algae in spring is nothing like so great as in the colder seas about Britain,
but it is most marked all the same. Rhizosolenia species follow Chaetoceras
species in the spring outburst in British Seas. They do the same here, although
the proportions of the species and the constitution of the catches are not identical
(see Text-fig. 4, p. 200).

Our spring maximum commences in August. From the point of view of the
calendar this would correspond with February in the Northern Hemisphere and
thus our phytoplankton maximum is relatively earlier than it is in British Seas.
In the latter it falls in March or as late as April. On the other hand, Steuer
records the diatom maximum as beginning in December, in the Adriatic at
Trieste, and continuing to the end of February. This is followed by another,
but weak maximum in June and July. This locality is, however, not so suitable
for comparison as an eceanic coast.

Evidently in the Northern Hemisphere as one passes south from the colder
waters of the north, the spring diatom maximum occurs a little earlier in the
year. Unfortunately plankton records are rather restricted from the point of
view of latitude. The latitude of Sydney corresponds roughly with the northern
latitude of Madeira, or with that of Los Angeles on the coast of California.

The temperature of the Irish Sea water varies between 7° C. winter and
14° to 15° C. summer, whilst Steuer records the surface waters off Trieste as
ranging from 5° C. winter to 25° C. in summer. The ocean water off Sydney is
perhaps slightly more equable, ranging from 16° C. winter to 23° C. summer; the
temperatures are distinctly higher than those of British Seas. It will be obvious,
therefore, that the spring diatom maximum commences at very different tempera-
tures in the regions named above, although there is the marked resemblance of
season.

Our diatom maximum in 1931 occurred just as the sea temperature was
beginning to rise after having reached the minimum for the year, but the total
increase in temperature during the first month of the diatom maximum was only
about 1° C. The spring diatom maximum in British Seas occurs at approximately
the same place in the temperature curve of the year as it does with us, the
actual temperature being, however, about 10° C. lower in British Seas. So far
it has not been possible to correlate definitely the initiation of the diatom outburst
with changes in sea temperature. The matter will be discussed later; the out-
standing fact to record at present is the existence of a diatom rhythm which
follows the seasons, and appears correlated with them on this South Pacific
coast in a manner similar to that discovered off the coasts of Europe and
recognized elsewhere in the seas of the Northern Hemisphere.

SCHIZOPHYCEAE.
A species of Oscillatoria has been recorded at odd times during the year.
The most remarkable Schizophycean, however, is the parasitic Richelia intra-
cellularis Schmidt, found parasitizing diatoms. We have found it for the first

196 MARINE PLANKTON OF THE COASTAL WATERS OF N. S. WALES, i,

time on the Australian coast (also the first record for the South Pacific) in a
large species of Rhizosolenia. This peculiar organism has now been recorded
from the Red Sea, the Indian Ocean, the Straits of Malacca, the Gulf of Siam
and the African Coast.

THE Protozoa.
Only those Protozoa which have appeared abundantly as constituents of our
plankton are recorded here.

MASTIGOPHORA.
Dinoflagellata.

The members of this group may well be taken first, for they should be
considered with the diatoms as belonging to the group of ‘producers’. Most
important are species of Peridinium, together with Ceratium furca. Ceratium
tripos is not uncommon with interesting oceanic and tropical forms as, for
example, the fascinating Ceratium ranipes Cl., C. palmatum Br. Schroeder, and
C. platycorne.

Species of Peridinium have been taken in every month of the year, the largest
catches being in November, 1931, on the same date as the largest catch of
Ceratium furca. Peridinium species are, on the whole, late spring and summer
forms, and this fits in with their general occurrence elsewhere. No attempt has
been made to separate the species in the enumerations, but the following have
been noticed: Pyrophagus horologium Stein., Peridinium ovatum (Pouchet)
Schutt., Diplopsalis lenticula Bergh., Gonyaulax spinifera Diesing.

Ceratium furca (Ehbg.) Cleve is found throughout the year, but is sparse
in the winter months from May onward until October (see Text-fig. 4). C. furca
is found over a wide range, absent perhaps from the colder waters of the Arctic
and Antarctic Seas. It is recorded from the North Atlantic, the South Atlantic,
the Indian Ocean, the Japanese Coast of Pacific, etce., viz., from the temperate
to tropical seas. The species occurs in the ocean plankton as well as that of
neritic waters.

Ceratium fusus Clap. and Lachm.—This species is of relatively rare occur-
rence here compared with the other species of Ceratium, and is of little significance
in the quantitative picture of the plankton during the seasons examined. It is
probably of cosmopolitan occurrence and of wider range even than C. furca.

Gymnodinium sp.—A species of Gymnodinium as yet undetermined, but prob-
ably new, is of special interest because of its extraordinary swarming habits in
the harbour of Port Jackson. It has been known to occur for years owing to
the fact that, when at its maximum, which lasts two or three weeks, it is
abundant enough to colour the harbour water red. For very many years
(actually since 1856) this has often been so striking an occurrence that letters
from irate correspondents have appeared in the Sydney papers asserting amongst
other explanations that blood from the abattoirs was being allowed to contaminate
the harbour. The attention of the public is still periodically aroused.

For three years we have observed this Peridinian appear in the month of
July or August, and the date has not varied more than three weeks in the
years 1930, 1931 and 1932.

Whitelegge examined the harbour water in 1891 when a similar red colora-
tion appeared in March and April. He wrote at some length on the organisms
present and concluded that the coloration of the water was due in the first case
to a new peridinian which he called Glenodinium rubrum and stated that later

BY W. J. DAKIN AND A. COLEFAX. 197

there was a marked prevalence of Gymnodinium spirale Bergh. At that time, and
for many years afterwards, there was no proper knowledge of the structure of
the unarmed dinoflagellates and it seems scarcely possible that Whitelegge’s
Glenodinium rubrum could be recognized to-day from the description given.
Again, from his illustration, it is certain that our Gymnodinium sp. which we
have found to occur so regularly in July and August is not the one he ealls
G. spirale Bergh.

There are other dinoflagellates which are responsible for red discoloration
of the sea, and it is possible that during the thirty years which have elapsed
and which have been accompanied by very considerable faunistic changes in
the waters of Port Jackson, other genera and species have taken the place of
earlier ones. In view of the absence of micropreparations and accurately detailed
drawings of Whitelegge’s specimens, it is impossible to go any further. The
greatest authority on the group, Professor Kofoid of the University of California,
has examined our species and he regards it as belonging to a new species of
Gymnodinium, but the structure is so delicate that more careful examination of
living material is now necessary before an exact diagnosis can be given.

The species never develops to the extraordinary state above described in the
outside waters, but it does occur in the ocean catches and often at later dates
than those noted for the harbour maximum.

There are other points of interest about this remarkable form. In the first
place, its maximum occurs when the water is at its coldest, whilst the Peridinians
are usually characterized by maxima in warmer seasons. In the second place, it
seems futile at present to try and correlate this extraordinary and regular ‘‘out-
burst” with chemical conditions. The only seasonal conditions which recur so
exactly are length of day (and consequently light) and possibly sea temperature.
One is reminded very forcibly that the organism itself may possess a very
significant inherent reproductive rhythm.

Dinophysis tripos Gourret.—The synonymy of this species is a very tangled
one, and is referred to at some length by Kofoid. In view of the difficulty of
nomenclature, it is not easy to discuss the matter of geographical distribution.
It is known from the Mediterranean Sea, the North Atlantic and South Atlantic
and possibly under the name D. caudata it is known from the Pacific. On the
other hand certain authors consider D. caudata a distinct species.

This flagellate is of common occurrence in our plankton catches and is present
throughout the year at odd times. The largest numbers and most regular presence
were noted in 1931 for the weeks between 3rd October and 1st November.

Noctiluca scintillans (Macartney), (Syn. N. miliaris Suriray)—Noctiluca has
only occurred three times in sufficient numbers to be noted on the counting plates
and these occasions were in October, December and January respectively. On
another occasion, however, in late summer, the author made a catch at night in
Sydney Harbour, well away from the open ocean and the haul was almost entirely
Noctiluca. Evidently the organism can at times be very common.

CILIATA.

A number of species of Tintinnidae have turned up in moderate numbers in
some of our catches, but the time of occurrence seems to bear little relation to
season. The following species have been recorded by us, all for the first time from
this part of the Pacific Ocean: Tintinnopsis vasculum Meunier, Tintinnopsis urnula
Meunier, Tintinnopsis radix (Imhof) Bdt., Favella campanula (Schmidt) Jorg.

198 MARINE PLANKTON OF THE COASTAL WATERS OF N. S. WALES, i,

(the two last mentioned are probably our most common species), Codonellopsis
ostenfeldtii (Schmidt) (of not infrequent occurrence on certain occasions),
Rhabdonella hebe (Cleve), Xystonella sp., Cozliella ampla (Jorgensen) Brandt.

There is evidence that these Tintinnids are much more common in the
estuaries than in the open ocean. One of us (W.J.D.) has been struck many
times by the abundance of Tintinnids in Australian estuarine waters. The same
prevalence has been noted by Mr. T. C. Roughley (verbal communieation) in the
Hawkesbury and George’s Rivers and Tintinnid tests have been found amongst
other organic remains in the stomachs of oysters. Amongst these we have
recognized from sketches, Codonellopsis ostenfeldi, Tintinnopsis sp., probably
urnula, and a Favella species.

RADIOLARIA.

The only common Radiolaria in our catches have been Acantharians like
Acanthometron, and colonial forms (without skeletons) like Collozoum inerme
Haeckel.

Acanthometron sp.—Whilst of frequent isolated occurrence the numbers only
became sufficient for enumeration on 25th October, 1931, 8th December, 1931, and
23rd February, 1932.

Collozoum inerme Haeckel.—Colonies were only numerous on one occasion,
in the month of April, although of sporadic occurrence during the summer months.

CoELENTERATA.

The most common coelenterates caught in our big nets or observed from
the boat have been:

ScypHozoa: Aurelia caerulea R.V.L., Pelagia panopyra Peron and Less.
SipHONoPHORA: Physalia physalis Linné. (? var. utriculus), Velella spirans Forskal,
Porpita porpita Linné, and Diphyes dispar Chamisso.

Hydroid medusae have been scarce, only Obelia medusae having played any
serious part in our catches. Probably the medusae of Hydroids would have been
more conspicuous had the catches been made closer inshore or in the sheltered
inlets where the adult forms lived.

Aurelia caerulea was particularly abundant in October, and in the same
month in 1931 and less so in 1932 great numbers of Pelagia panopyra entered
the harbour.

At the beginning of October, 1932, a great invasion of Physalia, Velella and
the Salp—Thalia democratica—came in from the north-east together with the
pelagic mollusc Janthina violacea. Millions of large Physalia could be observed
covering the sea surface, and later on the sea beaches. Bathers were kept out
of the water altogether on one day by the prevalence of these stinging creatures.
It is said that such an invasion has only been noted two or three times in the
past ten years.

CTENOPHORA.

Ctenophores are at times particularly common in the plankton. On one
occasion in the summer, Ctenophores were so abundant that a dredge (prawn-
mesh netting) became full as it was lowered to the bottom where it caught nothing
else. A broken mass of jelly two feet high was turned out of it on to the deck!

Amongst the common species is a Beroe sp., Bolina chuni, and a new species
of which a rather beautiful photograph is shown (PI. vii, fig. 1).

BY W. J. DAKIN AND A. COLEIAX. 199

PELAGIC MOLLUSCA.
STREPTONEURA.
Platypoda.
lanthina violacea Bolten.—An invasion of large numbers of this gastropod
occurred with Physalia, ete., during October, 1932.

Heteropoda.

Pterotrachea subgenus Huryops sp.?—On two or three occasions each year
(generally with an invasion of Salps and Siphonophores), heteropods have been
frequent in our catches with the big coarse nets. The most common species is a
member of the subgenus Huryops of Pterotrachea. It has not been possible as
yet to diagnose the species.

Firoloida kowalewskyi (?) Vayssiere.—Less frequently occurring than
Pterotrachea sp. is a species of Firoloida, very probably kowalewskyi, but females
are necessary for its correct determination, and so far only males have been
found; these agree perfectly with the diagnosis of the species. F. kowalewskyi
has been recorded from the Indo-Pacific region and the Atlantic.

Atlanta sp.—On two occasions small specimens of a species of Atlanta have
been common.

Pteropoda.

Creseis virgula Rang. var. conica.—This is the most common species of our
planktonic pteropods. It occurs sporadically throughout the year, and is found
most abundantly in catches of May, July, November, December and February.
This species is a cosmopolitan form, more characteristic of the warmest parts
of the great oceans. It has been recorded already off the Hast Australian Coast.

CHAETOGNATHA.

Specimens of Sagitta, mainly of small size, are ot fairly general occurrence
throughout the year. Several species occur and, up to date, these have not been
worked out. Sagittae have been most abundant in the summer months and
autumn, the poorest hauls so far have been those made in October, November
and early December.

CRUSTACEA.
Phyllopoda.

The Phyllopoda play a very considerable part quantitatively in the com-
position of the zooplankton of the locality investigated. Their seasonal distribution
is also most marked.

Hvadne nordmanni Lovén.—This species appears in the plankton after the
spring diatom maximum, in October (although its isolated occurrence before this
date is noted). It attained a maximum (in 1931) in November, and then gradually
fell off in numbers, to be followed by the other two species of the genus, which
seem rather more important in these waters. EH. nordmanni has a wide oceanic
range. It is found in British Seas and the North Atlantic, but during Hensen’s
plankton expedition was only captured north of latitude 40° in the Atlantic.

Evadne spinifera P. E. Miiller.—This species, after being absent throughout
the winter of 1931, appeared after the spring maximum of the phytoplankton in
the early summer months and continued practically throughout the summer. The
largest numbers recorded in 1931 were in January. The numbers fell off in
March and the species practically disappeared in May.

The species has a wide oceanic distribution. It is common in the Mediter-
ranean and the North Atlantic, with a tendency to be a southern form (Sargasso

200 MARINE PLANKTON OF THE COASTAL WATERS OF N. S. WALES, i,

Sea) there. It is recorded from the Indian Ocean and the west coast of
Australia. Kramer states that he found it off the Australian Coast, but does
not list it with the other species. It has been recorded by the Scottish National
Antarctic Expedition from one station in the South Atlantic, but that was in
Latitude 28° 8’ and not in the Antarctic.

EHvadne tergestina Claus.—This cladoceran is the most ubiquitous with us,
occurring at odd times in the winter. Its maximum developed (in 1931) a little
after that of spinifera, late in December, and the species continued abundant until
May. :

The distribution of HL. tergestina is wide and the species has been recorded
from the north-west Atlantic Coasts, to one station by the Scottish Antarctic
expedition in the South Atlantic, and from Fremantle, Western Australia. Kramer
records an Hvadne from Auckland, New Zealand, as a new species—#H. aspinosa
and also from Jervis Bay, but it is supposed by Hansen that this was a mistaken
identification and that the species is really #. tergestina. This is now almost
certainly proven for, whilst H. tergestina is common with us, no other fvadne
species has been recognized apart from those here recorded.

Jan. Feb Mar. Apr. MAy Jun. TuL Aue Sep. Ocl Nox Dee Jan. Feb,
Asterionella japonica ne Seen

Chaeioceras sf.
Razosolenia sp.
Slebhanopyas Sp
Thalassrosira sp.
Ceralium furea

Acarha clavs¢
Faracalanus ha
Temora sp.
Evadne sf.
Fedon sp.

Penilia schmacher.

Text-fig. 4—Times of maxima of the chief diatoms and dinoflagellates of
the New South Wales coastal plankton.

Text-fig. 5.—Times of maxima of the chief Copepoda and Cladocera of
the New South Wales coastal plankton.

Podon polyphemoides Leuck.—This species is far less abundant than the
Evadne species and its temporal distribution more limited. After a complete
absence during the winter, it appeared in 1931 late in December at the same time as
E. tergestina. It rapidly attained its maximum and seems almost to have dis-
appeared again in the month of January, 1932. The same thing occurred at the
beginning of January, 1931.

The distribution of Podon polyphemoides, like that of the other cladocerans
mentioned, is very wide. It has been recorded from the North Sea, North
Atlantic, Mediterranean, South Atlantic, and at Auckland (New Zealand). The
latter reference is particularly interesting, as it brings it for the first time into
the South Pacific.

BY W. J. DAKIN AND A. COLEFAX. 20)

Penilia schmdckeri Poppe.—This large and handsome cladoceran is quite
common in our catches during certain seasons. It occurs in odd numbers during
the winter, but its definite maximum begins only in late December (1931), and
it then continues numerous throughout the summer and autumn. In 1921 it
became rare in June, so that its season appears to be as long as and to coincide
with that of Hvadne tergestina.,

The distribution of this species is curiously wide within certain limitations,
but it has already been recorded from Hong’ Kong, from the harbour of Vera
Cruz, from the Gulf of Guinea (Atlantic) and lastly by Kramer from Auckland
and Port Jackson. It is not recorded in the Nordisches Plankton as occurring in
the North Atlantic or adjacent waters. Its distribution is, therefore, rather
different from that of the other Cladocera.

Summary ReGaArpDING DISTRIBUTION oF CLADOCERA.

The Cladocera are represented by five common species whose seasonal distri-
bution in our waters in the period 1931-32 is indicated approximately in Text-
figure 5.

OSTRACODA.

The only ostracod which has played any part worthy of note in our catches
is Huconchoecia aculeata Scott, and the numbers of this have been too small to
record. It would have been necessary to count the examples in whole catches.

CoPpEPODA.

From the importance of this group as an economic unit in the plankton, the
attention paid to it has far exceeded all other groups. It was necessary, indeed,
before conducting any serious statistical studies, to be able to recognize the
different species of copepods. Since there are no papers on the plankton of
this region, a rather complete taxonomic study has thus been necessary, even
whilst confining attention to the more common species.

From the North Atlantic waters and the waters washing the coasts of
Europe, more than 300 species are recorded in the “Nordisches Plankton’’, but
out of this number, only seven are enumerated as conspicuous plankton constituents
in Lohmann’s very full studies, and only eight in Herdman’s lists. So far we
have recorded only about 110 species, but it must be remembered that we have
not been specially seeking them. Of this number, about ten can be considered
as items of regular and significant occurrence, but time has already shown that
probably some of the larger copepods have not been captured in the special
quantitative nets, and in future silk nets of greater size and coarser mesh will
be dragged more rapidly for them. They are undoubtedly sought and captured
by fish.

It is evident that the species of Copepoda which play a big part quantitatively
in the plankton, both here and in Huropean Seas, are few in number, but these
species occur in incalculable myriads.

Like the diatoms and the Cladocera, the copepods which are recorded below
have generally a remarkably wide range in the world’s seas, and it is very
striking to find that the two most common species in this locality also belong
to the list of the eight most common copepods of the Irish Sea. They are
Acartia clausi and Paracalanus parvus, and it would probably be not incorrect
to say that these were the two commonest crustaceans in the world’s seas, indeed
possibly the two commonest of all aquatic metazoa.

202 MARINE PLANKTON OF THE COASTAL WATERS OF N. S. WALES, i,

The following is a list of the species discovered by us in the plankton of the
coast near Sydney. Most of these species are recorded for the first time in these
waters; a few copepods are recorded in Whitelegge’s list of the Invertebrate
Fauna of Port Jackson, but these are taken from the records of the odd catches
made by the “Challenger”? in 1874, and for some reason are by no means repre-
sentative of the copepod fauna as it occurs to-day. Recently, however, an account
has been published of the Copepoda of the “Terra Nova” Expedition (Farran), and
since this expedition’s track crossed the Pacific between New Zealand and
Australia, we have a valuable record of a considerable number of South Pacific
copepods from a locality not so very far away. The seasonal distribution of the
more important types is shown numerically in Table II, and of the two most
common species in Text-figure 5.

Text-fig. 6.—Some of the more important diatom species of the New South
Wales coastal plankton. 1.—Planktoniella sol (Wallich). 2.—Chaetoceras
didymum Ehr. 3.—C. sociales Lauder. 4.—Rhizosolenia’ alata Brightw.
5.—R. stolterfothii H. Perag. 6.—R. sp. 7.—Dityllium Brightwelli (West).
8.—Hucampia zoodiacus EHhr. 9.—Leptocylindrus danicus Cleve. 10.—
Streptotheca thamensis Shrubs. 11.—Asterionella japonica Cleve and Moller.
12.—Climacodium frauenfeldtii Grun. 13.—Rhizosolenia sp. with the parasitic
Schizophycean Richelia intercellularis Schmidt. 14.—Thalassiosira rotula
Meunier.

BY W. J. DAKIN AND A. COLEFAX. 203

Calanidae.

Calanus brevicornis.—South Atlantic, Southern Ocean, Mediterranean, off New
Guinea. Not recorded in New Zealand waters by “Terra Nova” Expedition. Rare
except for one big catch in September. It is, however, the most common copepod
in the stomach of the fish, Apogonops, caught in October, off Newcastle.

Calanus darwiniit (Lubb.) = Undina darwinii.—Common off New Zealand,
Tropical Atlantic, not recorded Northern Atlantic?. Apparently not recorded
North Pacific off California. Recorded from South Pacific (“Siboga’ Expedition)
and off Australia (‘Challenger’ Expedition). It was present in our catches in
April and May.

Calanus jfinmarchicus (Gunner).—Cosmopolitan, North Pacific, North
Atlantic, Arctic, Tropical Atlantic, common New Zealand waters and in Melbourne
Harbour. The Australasian Antarctic Expedition collected the species south of
Tasmania. This species occurred infrequently but at odd times throughout the
year.

Calanus gracilis Dana.—Cosmopolitan, North Atlantic, Tropical Atlantic,
Indian Ocean, Pacific Ocean, Mediterranean, North Pacific off San Diego, rare.

Calanus minor Claus.—Tropical Atlantic and subtropical, Southern Pacific
off New Zealand, North Pacific off California. This species is the most abundant
of the Calanus in our catches, but only reached figures which justified counting
towards the end of March, 1931.

The species is closely related to Calanus finmarchicus, from which it differs,
amongst other things, in having the head and first thoracic segment fused. We
have had the typical C. minor in our catches, but certain specimens have been
captured in which the head and first thoracic segment are separate, although other
characters are of the C. minor type. Other features are present of intermediate
character.

Text-fig. 7—Common Tintinnids of the New South Wales coastal plankton.

1.—Tintinnopsis radix (Imhof) Bat. 2.—Codonellopsis ostenfeldti
(Schmidt). 3.—Favella campanula (Schmidt) Jérg. 4.—Rhabdonella hebe
(Cleve).

Calanus patagoniensis (Brady).—Tropical Atlantic. Not recorded in ‘“Siboga’”
Report, nor in ‘‘Terra Nova” Report from South Pacific. This species only turned
up twice in our catches, those of 9th August, 1931, and in the same month of
1929.

204 MARINE PLANKTON OF THE COASTAL WATERS OF N. S. WALES, i,

Calanus pauper Giesbr.—Not recorded off Coast of California. Farran recorded
one specimen only off New Zealand, but this species, with Calanus minor, is the
most abundant species of the genus in our waters. It occurred chiefly in the
summer months of 1931, more particularly in January and February.

Calanus tenuicornis Dana.—North Atlantic, North Pacific, Mediterranean Sea,
South Pacific and common off North Island, New Zealand. This has not been
common in our catches, but its abundant presence in our waters was indicated
by the stomach contents of the fish Apogonops.

Calanus vulgaris (Dana).—North Atlantic, Tropical Atlantic, South Atlantic,
off Australian Coast (‘“Challenger” Expedition). Not uncommon in our catches,
but most specimens have occurred between March and May.

Hucalanidae.

EHucalanus attenuatus Dana.—A species of cosmopolitan range and recorded
off the east coast of Australia by ‘Challenger’ Expedition.

Hucalanus crassus Giesbrecht.—Recorded from the Atlantic, Pacific and Indian
Oceans. Odd specimens were taken by us in April and May.

Hucalanus elongatus (Dana).—This species is recorded as common off New
Zealand by Farran. It occurred in our catches at an immature stage.

Hucalanus monachus Giesbrecht.—This species was represented in our catches
by odd specimens in March, May and August.

Rhincalanus cornutus (Dana).—Farran found this twice off New Zealand. It
had previously been recorded from the Atlantic, Indian and Pacific Oceans. It
occurred twice only, in July and November.

Rhincalanus nasutus Giesbrecht—This is common off North-west New
Zealand according to Farran. It is a species of world-wide distribution, but never
eommon with us. It was most frequent in May and August.

Mecynocera clausi I. C. Thompson.—Although this form has never been
numerically strong, several individuals are usually taken in every haul, being
readily recognizable by the enormously elongate antennae.

Paracalanidae.

Paracalanus aculeatus Giesbrecht.—This species, which is rare compared with
P. parvus and of restricted range, was recorded once in the summer months. It
is stated by Farran to be frequent in the tropical Atlantic but was captured once
by the “Terra Nova” off New Zealand.

Paracalanus parvus (Claus).—This Copepod which is of very wide distribution,
being recorded from the Atlantic Ocean, North Sea, Black Sea, Red Sea, Indian
Ocean, and Pacific Ocean, is one of the two most common crustacea in our
plankton. It is quite extraordinary, therefore, that this species was not recorded
by the “Challenger” Expedition off our coasts. The Australasian Antarctic
Expedition captured it frequently in the far south. It occurs throughout the
year and is common at all times except during the winter months of August and
September. It develops an abundant maximum following the phytoplankton
maximum during late autumn (May and June). It is noteworthy that this
copepod is one of the six or seven species which are so abundant in Northern
Seas as to be generally found in numerical estimations.

Acrocalanus gibber Giesbrecht.—This species was originally recorded from
the East Pacific Ocean by Giesbrecht. It is known also from the Red Sea. It is
not recorded in the ‘“‘Terra Nova” catches either from the Atlantic or Pacific. The

BY W. J. DAKIN AND A. COLEFAX. 205

species is rare in our catches, but turned up in April, 1931, in sufficient numbers
to be counted.

Acrocalanus gracilis Giesbrecht.—This species has been of frequent occurrence.
It was conspicuous in January, 1931, and present during March, April, May and
August of that year. It also turned up in January and February, 1932. The
range of the species appears very limited. It was first recorded from the East
Pacific.

Calocalanus plumulosus (Cls.).—This species has been recorded as plentiful
off New Zealand. It occurs in the Atlantic Ocean and Mediterranean Sea. It has
only been found once in our plankton catches.

Pseudocalanidae.

Ctenocalanus vanus Giesbr.—This species has been recorded from the Mediter-
ranean, Atlantic Ocean, Pacific Ocean and the Antarctic, and Farran remarks that
it is one of the most plentiful species in tow-nettings taken under the ice (‘‘Terra
Nova’). It has provided us with a demonstration of the probable deficiency of
plankton-catching weapons for, although of characteristic appearance and easy
recognition, only one solitary specimen has been obtained by us.

Clausocalanus furcatus Brady; Clausocalanus arcuicornis Dana.—These have
‘been two common constituents of our catches and although they are never so
abundant as Paracalanus parvus, they may be ranked as members of the small
group of really common copepods in these waters.

Scolecitricidae.

Scolecithrix danae (Lubb.).—Recorded from the Mediterranean, Atlantic,
Indian and Pacific Oceans and regarded as a more or less tropical form, this
species is noted as common off North Island, New Zealand. It was only recorded
in July in our plankton.

Centropagidae.

Centropages bradyi Wheeler.—Recorded from the North Atlantic by Wheeler
and from several stations off New Zealand by Farran. Its only occurrence in our
plankton was on 5th March, 1930.

Centropages calaninus (Dana).—Recorded from the Malay Archipelago and
from the Indian Ocean. This species occurred in considerable numbers in the
eatches of 29th March, 1931, but that is the only date.

Centropages chierchiae Giesbrecht.—This Centropages is of wide range, already
recorded from the more tropical Atlantic Ocean and the Indian Ocean. Jt was
recorded once from our plankton of December, 1929.

Ceniropages furcatus (Dana).—This species, according to A. Scott, has a
wide range in the tropical seas. It was the most abundant species collected by
the “Siboga’ Expedition in the Malay Archipelago. It is the only Centropages
’ species recorded by the “Challenger” from our own locality. The species was
relatively common on 2nd February and 7th April, 1931. It also occurred
sparingly in May and June.

Centropages orsinii Giesbrecht.—This species of Centropages which is an
Indian Ocean form, recorded also from Red Sea and Malay Straits, is our com-
monest species of the genus. This fs rather a surprising feature, for there appear
to be no records indicating a wider distribution in the Pacific and the species is
not recorded from New Zealand waters by Farran. The species is most abundant
in late summer; specimens were common from January to May.

G

206 MARINE PLANKTON OF THE COASTAL WATERS OF N. S. WALES, i,

Centropages violaceus (Claus).—Recorded from the tropical Atlantic, and
parts of the temperate Atlantic, Mediterranean Sea, and, according to Farran,
from the more northerly of the New Zealand stations. Of rare occurrence in our
catches—recorded in September.

Pontellidae.

Pontellopsis regalis (Dana).—Recorded from Atlantic, Indian and Pacific
Oceans. Rare in our plankton. Recorded in one catch, 10th April, 1931.

Labidocera acuta (Dana).—Recorded from the tropical waters of the Atlantic,
Pacific and Indian Oceans, and collected by the “Challenger” off our coasts. Not
recorded, however, from New Zealand stations by Farran. This species is of very
common occurrence in the New South Wales plankton. Specimens have been
taken in every month from January to August.

Labidocera acutifrons (Dana).—Recorded from the tropical Atlantic and
Pacific Oceans and collected by the “Challenger” off our coasts. Much more rare
than L. acuta and only found in April, 1930.

Labidocera brunescens Czerniawsky.—Recorded from the Atlantic and Indian
Oceans. Taken once by us in January, 1930.

Labidocera cervi Kramer.—This is the only copepod species limited to the
waters between New Zealand and Australia, which plays any important part in
our plankton. It was discovered by Kramer, who only found one specimen, a
female, in the Hauraki Gulf, New Zealand, in 1893. Six years later it was found
again by Brady in copepoda from New Zealand. Farran discovered it in a few
of the “Terra Nova’ catches from New Zealand waters and a variety in Melbourne
Harbour. We are apparently the first to find it really abundant. It was common
in our catches during the summer and in numbers sufficiently great to be
counted in small samples in April, 1931.

It might be pointed out that the males in our collection differ slightly from
the specimen figured by Farran in details of the fifth feet. The variation is slight
and may be a local racial difference.

Labidocera detruncata (Dana).—Recorded only from the Indian and Pacific
Oceans. The species was only recorded in the catches of 1st November, 1931.

Labidocera Kroyeri (Brady).—Recorded from North Atlantic, Pacific and
Indian Oceans and Malay Siraits. A few specimens were present in April in 1930
and 1931.

Candacidae.

Candacia bipinnata Giesbrecht.—Recorded from the tropical Atlantic and the
Pacific Ocean (North and South), Malay Straits, etc. This species was the most
abundant according to Farran in New Zealand waters. It occurred in our catches
in April and May, but probably is more common in our waters.

Candacia ethiopica (Dana).—Recorded chiefly from tropical seas, Atlantic,
Pacific and Indian Oceans, Malay Straits and off New Zealand. Occurred not
infrequently, particularly in April and May.

Candacia pachydactyla Dana.—Recorded from tropical Atlantic, Pacific Ocean
and Indian Ocean. The species was present on one day in April, 1930.

Candacia pectinata Brady.—Recorded from off Port Jackson by the ‘“‘Challenger’’
Expedition, and from the North Atlantic, Mediterranean Sea, and Indian Ocean.

The occurrence of this species is fairly common, which is striking in view
of its recognition amongst the few copepods captured by the ‘“Challenger’’
Expedition in our waters and the absence of the name both in Farran’s list from

BY W. J. DAKIN AND A. COLEFAX. 207

New Zealand waters and the “Siboga” list of Malay Strait forms. It was most
common in a night catch taken in November, 1931.

Candacia truncata (Dana).—Recorded from the Atlantic, Indian and Pacific
Oceans; was present in our catches of May, 1931.

Temoridae.

Temora discaudata Giesbrecht.—This is practically a Pacific Ocean copepod,
although it has been recorded from the Red Sea. Most records are from tropical
waters. It is not recorded from New Zealand waters.

The species is of rare occurrence in our catches, but has been noted in
several months, April, May, September, and most common in February, 1932,
when it was sufficiently abundant to be enumerated.

Temora stylifera (Dana).—Recorded from the warmer waters of the North
and South Atlantic Oceans and found commonly in the “Terra Nova” catches from
the tropical Atlantic. Rare in our catches except in February, 1932.

Temora turbinata (Dana).—This copepod species is quantitatively one of
the most important in our plankton. It is a well known Pacific Ocean form,
recorded as of great abundance off New Zealand and common in the Malay Straits.
T. turbinata occurs throughout the year in our seas; it presented one maximum in
the year 1931-32 during the summer months with the peak in February.

Acartiidae.

Acartia clausi Giesbrecht.—This copepod species shares the honour with
Paracalanus parvus of being the most abundant constituent of the plankton
crustacea. It is common throughout the year, but rises to two maxima, the
greater in October-November, and a lesser in January, although it is possible that
these two are parts of one. It will require further years of observation to become
familiar with the variations in seasonal distribution which are bound to occur.

This copepod has been recorded from both the cold regions and the tropics,
inorth and South Atlantic, the Indian Ocean, South Pacific and far down towards
the ice. It was not recorded from the New South Wales Coast by the ‘‘Challenger”’
and it does not appear in the Malay Straits collections of the “Siboga” Expedition.

Acartia centrura Giesbrecht.—A specimen of Acartia which seemed to resemble
A. centrura most closely was recorded in May, 1931. The species has been
recorded from the Red Sea.

Acartia danae Giesbrecht.—This species, recorded from the Atlantic and
Pacific Oceans, was recorded as rare in Malay Straits. It is common off New
Zealand. It is rare in our catches, being recorded only on ist May, 1931, and in
January, 1930.

Tortanidae.
Tortanus barbatus (Brady).—Recorded Pacific Ocean (Philippines), Malay
Archipelago. Occurred on one occasion, 1st November, 1931.

Oithonidae.

Oithona atlantica Farran.—Recorded from west and south-west coast of Ireland,
Southern Atlantic and Indian Ocean. This species has not been frequently
recorded. It is difficult to separate, when counting, from O. tenwis, but there is
no doubt that it is not at all common.

Oithona nana Giesbrecht.—Practically cosmopolitan and very common in
some regions. Only occurred in our catches on one day of October, 1931.

208 MARINE PLANKTON OF THE COASTAL WATERS OF N. S. WALES, i,

Oithona oculata Farran.—Recorded by Farran from Indian Ocean catches
also from near Samoa. Its occurrence in our plankton was sufficiently common
to enumerate in April and July, 1931; it was also present in August and January.

Oithona plumifera Baird.—Of wide range. Recorded from polar seas, Atlantic
Ocean, Indian Ocean, Pacific Ocean and common off New Zealand and in Malay
Straits. The species occurred at odd times throughout the year in our plankton
catches, but was most numerous in May, June, November and February,

Oithona setigera Dana.—Recorded from the Pacific, Indian and Atlantic
Oceans, Red Sea and Mediterranean, New Zealand Waters. Not uncommon in
certain catches in months of January, February, April and July.

Oithona tenuis Rosendorn.—This Oithona species which appears seldom in the
literature is apparently our most common species of the genus. It occurs at odd
times throughout the year (reaching counting intensity from June to August,
1931, and in November), and was present fairly regularly from December to
February. Two specimens were recorded by Farran in New Zealand waters. It
was previously known from tropical Indian Ocean and Atlantic.

Hctinosomidae.

_ Microsetella rosea (Dana).—Already recorded from the Atlantic, Red Sea,
Indian Ocean and Pacific Ocean. In the Nordisches Plankton only the female is
described and the male said to be unknown. Farran records it “a few times off
New Zealand’, but only gives dimensions of the female, so that it appears doubtful
whether the male has ever been previously seen. The species is quite common in
our plankton and was particularly abundant from January to end of May, with a
peak in March (1931). Nauplii were obtained in June and July. In view of the
non-appearance of males elsewhere in catches, it is particularly interesting that
many males were examined, and on 4th January, 1931, when 560 were counted
in a sample, the males amounted to about one-third of the total. A description
of the male will be given in another paper.

Monstrillidae.
Thaumaleus thompsoni Giesbrecht.—North Atlantic. Recorded on one
occasion, March, 1930.

Macrosetellidae.

Setella gracilis Dana.—Recorded from Atlantic, Pacific and Indian Oceans
and taken in small numbers off the New Zealand Coast. Recorded by us on one
day only, 3rd October, 1931.

Tachidiidae.

Huterpe acutifrons (Dana).—Recorded from North Sea, North Atlantic Coasts,
Red Sea, Indian Ocean, Pacific off New Zealand, especially Bay of Islands. This
is a very common copepod of our plankton. It was taken in almost every month
of the year and was particularly common in the winter months, June to September.
The biggest haul occurred in a night catch, so that probably the species is more
abundant in deeper water than we regularly ‘“fished’’.

; Oncaeidae.

Oncaea media Giesbrecht.—Recorded from the Mediterranean, Atlantic Ocean,
Pacific Ocean, Red Sea, Malay Archipelago, off New Zealand, and possibly down
to Antarctic ice.

Oncaea venusta Philippi.—Mediterranean, Atlantic Ocean, Pacific and Jndian
Oceans, common off New Zealand and Malay Archipelago. This copepod species
is quite common in our catches and occurs throughout the year. The greatest
abundance occurred in April and May, 1931.

BY W. J. DAKIN AND A. COLEIFAX. 209

Corycaedidae.

Corycaeus clausi F. Dahl.—Recorded from Sargasso Sea and warm Atlantic,
Mediterranean, doubtful from Pacific. This species occurred on two occasions
only, but was not uncommon on Ist May, 1931.

Corycaeus crassiusculus Dana.—Recorded doubtfully from Atlantic, an Indo-
Pacific species. Common off New Zealand Coast. Recorded at odd times during
the year (January, February, April and May).

Corycaeus speciosus Dana.—Recorded from Atlantic, Pacific and Indian
Oceans; recorded as scarce off New Zealand. Only found once in our catches
(May, 1931).

Corycaeus (Corycella) carinatus (Giesbrecht).—Atlantic, Indian and Pacific
Oceans (tropical part). Common in our plankton from April to June, 1931.

Corycaeus (Corycella) concinnus (Dana).—Pacifie Ocean (tropical south-west
region), and tropical part of Indian Ocean. This copepod species was very
common in our catches between January and June, 1931, particularly on 20th
January and 10th April.

Corycaeus (Corycella) gibbulus Giesbrecht.—Recorded from Indian Ocean and
Pacific Ocean. Occurred only once in our plankton on 5th March, 1930.

Corycaeus (Corycella) rostratus Claus.—Recorded from Atlantic, South
Pacific and Mediterranean Seas. Was not uncommon in our plankton in January,
1932, but occurred also in September and November and December, 1931.

Corycaeus sp.?—A corycaeid appeared in our catches with considerable regu-
larity, which is not only different from the species hitherto described, but which
appears to invalidate the separation of the genus Corycaeus and the genus
Corycella in the scheme adopted by Dahl.

Corycaeus (Ditrichocorycaeus) africanus F. Dahl.—Recorded from the African
Coast. The species occurred infrequently and chiefly in catches taken in January
and April, 1931, also in May, June and August.

Corycaeus (Ditrichocorycaeus) andrewsi Farran.—Recorded from Christmas
Island, West Coast of Ceylon, and off Sumatra and New Guinea. The species
occurred on one date only, 3rd October, 1931.

Corycaeus (Ditrichocorycaeus) tenuis Giesbrecht—Recorded from eastern
part of tropical Pacific and West Pacific. Occurred in our catches on one date only,
ist May, 1931.

Corycaeus (Onychocorycaeus) catus F. Dahl.—Recorded from Indian Ocean
and Pacific Ocean. This is not uncommon in our plankton catches and was
counted during the period February to July, when it was continually present,
reaching a peak in February and late April.

Corycaeus (Onychocorycaeus) giesbrechti F. Dahl.—Recorded from Mediter-
ranean, Bermuda and Pacific Ocean. Occurred only once in our plankton catches
on 2nd August, 1931.

Corycaeus (Urocorycaeus) furcifer Claus.—Recorded from Mediterranean,
Atlantic Ocean, East Pacific and Indian Oceans. Occurred on odd occasions rarely
in our plankton.

Sapphirinidae.

Sapphirina nigromaculata Claus.—Of world-wide occurrence; Atlantic and
Indian Oceans, Pacifie Ocean and Malay Archipelago, but not recorded by Farran
in New Zealand catches. Recorded twice in our plankton.

210 MARINE PLANKTON OF THE COASTAL WATERS OF N. S. WALES, i,

Sapphirina angusta Dana.—Of world-wide distribution. Recorded from New
Zealand waters. Recorded on one day in November, 1930.

Sapphirina opalina Dana.—Of world-wide distribution. Recorded from New
Zealand waters. Recorded on one day—18th July, 1931.

Copilia mirabilis Dana.—Recorded from the Atlantic Ocean, Indian Ocean and
Pacific Ocean. The species was recognized on one occasion in our catches, 21st
June, 1931.

SCHIZOPODA.

Nyctiphanes australis—Apart from the Copepoda, this is perhaps the most
abundant Crustacean in our plankton, and certainly one of great importance in
the food chain of certain of the fishes. Our normal series of catches does not
contain many adults, presumably due to insufficient speed of towing when making
the hauls, but larvae in various stages of development have been taken, particu-
larly in late winter and early spring. However, catches made from the more
rapidly moving trawlers have proved more successful, both adults and larvae
coming up.

N. australis occurs in such enormous numbers off the New South Wales coast
that at times the sea is coloured blood-red, and on one occasion (Aug., 1930), a
solid mass of some two litres volume was taken in the small coarse plankton
net after a three-minute haul from the stern of a trawler. The red patches
marking the presence of Nyctiphanes were some six feet square, and were passed
by the boat every few seconds. They were accompanied by numbers of fish larvae
and the copepod Sapphirina. The solid mass of Nyctiphanes, when emptied from
the net, emitted a beautiful blue-green phosphorescence.

In March, 1931, an almost pure catch of the larvae was taken from a trawler
by one of us (A.N.C.) off the coast of New South Wales near Merimbula, whilst
early and late Calyptopi have occurred in the normal series of catches from the
station off Sydney Heads. We have also found large quantities of Nyctiphanes
in the stomachs of tiger flathead (Neoplatycephalus macrodon Ogilby), whilst the
larvae and occasional adults form part of the food of the fish Apogonops, which
is, in turn, preyed upon extensively by the tiger flathead.

UROCHORDATA.

The Urochordata have played no inconsiderable part in our catches, species
of Oikopleura being particularly abundant over a long period of each year. At odd
times the coastal waters are invaded by great swarms of Salps and then the nets
fail to give true pictures of the rest of the plankton, for the nets are soon
rendered almost impervious to water by the clogging jelly of the Salps.

Oikopleura sp.—Species of Oikopleura were present throughout the year and,
whilst high numbers were reached in summer, there were occasions with almost
equally high figures in August.

Like the neighbouring genus, Fritillaria, Oikopleuwra occurs in enormous
numbers at times. In October, 1930, an almost pure catch was taken from the
deck of a trawler off Botany (near Sydney). They were present in such quantity
that the sea was coloured bright orange and had the appearance of being
sprinkled with millions of minute rice grains; the organisms were confined to
a belt of water some twenty feet wide, marking, presumably, the course of a
minor ocean current.

It so happened that one of us (W.J.D.) was at the Sydney Heads Station
at the same hour, also for the purpose of making plankton catches, but here

BY W. J. DAKIN AND A. COLEFAX. 211

a very different result was obtained; the small coarse plankton net was filled
practically solid with small Salps (Thalia democratica).

It was also noticed that the sea was teeming with Coelenterates (principally
Siphonophores), whilst the copepod Sapphirina was so abundant that the separate
individuals could be detected in the water in the daytime, by reason of the
iridescent flashing of the cuticle.

Fritillaria pellucida Busch.—F'ritillaria species are far less frequent than
Oikopleura. The species listed here was abundant from February to April, 1931,
and again about the latter part of August, and in October and November. On one
occasion, however, on 3rd October, 1931, the deep net caught 40,000 whilst prac-
tically none were present in the surface waters. The highest catch otherwise,
was about 2,000.

The species is known from all the warm seas, and is noted rarely off the
Californian Coast in the Pacific, where apparently it was only recorded during
the summer. However, in the Mediterranean, it has been found abundantly in
the winter months.

Salpa fusiformis Cuvier.—Only the aggregated zooid form has been obtained
—in October and December (fairly common). The species has been recorded over
a wide range.

Thalia democratica (Forskal).—This is the common Salp of New South Wales
waters. Its appearance is sporadic, but when present it often occurs in very
large numbers. Its chief appearances have been in September, October and
November, with smaller numbers during the summer. Both the aggregate and
solitary forms occur.

The species has the widest recorded distribution of all the Salps.

Pegea confederata (Forsk&al).—One specimen of this large Salp was obtained
in a catch in December, 1931. It was the aggregate form. The species has been
recorded for the warm waters of the Pacific and Atlantic. The solitary form of
this Salp is unusually rare in the world’s collections compared with the aggregate
form.

Doliolum sp.?—Doliolum is a rare visitor compared with the Salps. Its most
abundant visitation was in August, 1931.

LARVAE.

Lamellibranch larvae—Common during the summer months.

Veliger larvae.—These occurred throughout the year, but were most common
in summer.

Ophioplutei.i—The Ophiopluteus larva of the brittle starfish occurred in the
catches from June to August and then at odd times until December. The maximum,
in 1931, was reached in July. It is as yet impossible to say to what species they
belong.

Actinotrocha.—The beautiful larvae of Phoronis occurred on one occasion in
the catch—seven or eight being present on 1st November, 1931.

Peneus.—The larvae of Penaeids occurred in our catches on several occasions
and this is the first time they appear to have been found in the sea round the
coast of Australia. In fact much mystery has shrouded the place of reproduction
of the Penaeid prawns, some species of which are extraordinarily abundant in
estuaries and coastal ‘lakes’ connected with the sea.

Mysid stages were present in March, April and May. Nauplii and eggs were
obtained from March to August, with a peak in May. Unfortunately, it is

212 MARINE PLANKTON OF THE COASTAL WATERS OF N. S. WALES, 1,

impossible to say from the eggs to which species they belong. Reference to
the breeding of the common species is being made in another place.

Cirripede nauplii.—Cirripede nauplii were present throughout the year and
with little fluctuation in abundance.

Ascidian larvae.—Ascidian eggs and larvae were abundant in some catches
in October and particularly abundant in certain inshore catches in which the
hatching of the eggs under the microscope was observed.

Other larvae.—Many different kinds of Crustacean and other larvae occurred
in our catches, but as casual occurrences, playing little part to speak of quanti-
tatively, although furnishing interesting material for future researches on life
histories.

FisH Eaes.

Fish eggs have at times been very abundant in our catches and we have
been, from the outset, most keen to collect this constituent of the plankton. The
number of different common kinds captured up to date, appears, however, to be
relatively small, and, in fact, may not be more than six. The earliest egg to
be recognized by us was the anchovy, which is very distinctive on account of
its elliptical shape and characteristically segmented yolk.

The most striking type of all, however, is one at present referred to for
convenience as a “blue” egg. Actually, there are at least two different kinds of
“blue” eggs according to measurements (one 1-6 mm. in diameter, and the other
1:3 mm.), but these share the common feature of a “shell” with a distinct blue
tinge. The perivitelline space is relatively large, the yolk is segmented and it
is considered, therefore, that these eggs may be clupeid (possibly those of
Sardinia neopilchardus). They have been found with early and late embryos,
whilst the very characteristic just-hatched larvae have also been secured. These
agree with a clupeid diagnosis.

In addition to the above, there are two other types of eggs which have been
common in our catches, but these, apart from their size, have not been marked
by any very special features, and it is hoped to treat them, together with the
rest, in greater detail in a future paper.

We are chiefly concerned in this survey with the abundance of the eggs in
the plankton during the period covered by the present communication.

High figures were reached in January, 1931, the ordinary coarse surface net
yielding 329, of which 112 were “Anchovy” eggs and the remainder undetermined.
On two other occasions in the same month, the catches were 279 and 561 eggs
respectively.

The number continued high (March 5, 644 in one net and 595 in another)
during the following months, but the Anchovy eggs had disappeared. The so-
called “blue” eggs (two kinds) appeared with startling abruptness in June (1070
in one net haul on 2ist June) and continued until August. The period June—
August represents therefore, the period of maximum of the blue eggs, although
some seem to occur at odd times throughout the year. Most of the “blue” eggs
have late embryos in them, early stages being rare.

After August the egg catches remained low until November, when the Anchovy
eggs commenced to appear. The Anchovy eggs continued to be present till
January, 1932, although not in large numbers during this last summer month.
Summarizing, we may say that in the period investigated, there were three
periods of abundance in regard to fish eggs, viz., (1) Anchovy eggs, November—

BY W. J. DAKIN AND A. COLEFAX. 213

January; (2) a period, January to April, marked probably by the occurrence
together of eggs of several unknown species at the same time and with a peak in
March; and (3) a winter maximum, between June and August, of the “blue” eggs.

GENERAL CONCLUSIONS.

The plankton at the station chosen, four miles out from the coast of New
South Wales in the latitude of Sydney, is more or less free from littoral forms.
It is a complex of oceanic and neritic species and, as one might expect, in view
of the fact that there is a tropical current setting southward in these Eastern
Pacific waters, one finds a conspicuous tropical and oceanic element, which is
particularly marked in the zooplankton.

On the whole, the larvae of benthoic forms are not strikingly abundant, but
this again may be explained by the situation of our station and the effort made
to avoid estuarine waters and close inshore waters. Night catches which were
made closer to the harbour entrance, presented a very different picture in this
regard, but these are not treated at all in this paper.

A distinct periodicity has been visible in connection with most of the
important constituents of the animal plankton (see Table II) and, whether it be
directly due, or not, to the fact that the phytoplankton (see Table I) shows a
definite maximum in spring, with another in late autumn, there is a clear
indication that the chief constituents of the zooplankton have their maxima about
the same times, and just after those of the phytoplankton. Thus the zooplankton
presents a maximum in the summer with peaks in the early summer (November
to December) and another in autumn. The zooplankton maxima referred to are
due to copepoda and cladocera. In addition, however, there are occasional
invasions of enormous shoals of Salps and Ctenophores, Siphonophora, and
Tanthina.

A remarkable feature of the plankton occurred during a fortnight in October,
1932, when the ocean surface was covered with millions of Physalia, Velella and
Iunthina. The invasion was so great that the dried bladders of Physalia and
Velella and the blue shells of Janthina strewed the. ocean beaches over a stretch
of many miles.

Curiously enough a letter in Nature of 29th October, 1932, reported an exactly
similar occurrence on the south-western shores of England and Ireland during
August.

Presumably the occurrence of these two unusual events (it is years since
an invasion of such an extent has been noted on our coast), at about the same
time is merely coincidence. It is difficult to regard it otherwise, seeing that we
are in the Southern Pacific and the British swarms come from the North Atlantic.
One might, however, wonder whether some particular cycle of circumstances had
led to an unusual development of the organisms in question in tropical ocean
waters, like the numeral fluctuations so well known in the case of more confined
terrestrial species of animals. But then it will be difficult to account for the
simultaneous abundance of species so far removed zoologically as Physalia and
Tanthina.

Perhaps, therefore, we had better think of it as just coincidence and look
for the explanation solely in a similar sequence of meteorological conditions in the
regions concerned. Certainly the weather off Sydney during the three months
preceding, and since the event has been somewhat unusual.

214 MARINE PLANKTON OF THE COASTAL WATERS OF N. S. WALES, i,

The State Meteorologist (Mr. Mares) has kindly sent me a comparison of the
winds during August-November, 1932, with those prevailing during this season
through the past 18 years. From this it would appear that sea breezes (i.e.,
winds with an easterly component) have been more frequent than usual and,
moreover, the velocities have been greater. It seems reasonable, therefore,
to suggest that there has been a greater tendency during the period in question
for an easterly surface drift to bring shorewards certain organisms of the tropical
southward-flowing current which are only infrequently stranded in such numbers.

We have referred to the occurrence of different species in this paper. There are,
of course, far more species in our plankton than enumerated or captured and,
if other modes of catching were adopted, some of these (chiefly microplanktonic
or nannoplankton forms) would no doubt be important also from the quantitative
‘point of view.

The seasonal distribution of the diatoms has already been described at the
end of the diatom section of this paper. Although many years of work will be
necessary before seasonal fluctuations are thoroughly understood, some additional
observations, made by one of us before our ocean station was established, are
sufficiently in agreement with the rest to warrant a little discussion on the
factors which may control the seasonal changes.

The actual demonstration of this seasonal fluctuation in the plankton produc-
tion of New South Wales waters emphasizes, however, the need for further
observations in other latitudes on the East Coast of Australia, and more particu-
larly to the south and in Tasmanian waters. Surface catches made with one
of our ordinary nets from trawlers 200 miles south of Sydney have often yielded
very different catches. We have never made huge catches of Nyctiphanes at
our station, but enormous numbers have been taken both north and south of
it, south in July and August, and north in October.

The total variation in the quantity of the zooplankton throughout the year
is not great. It might play little part in the migrations of the particular species
of fish which are just at present of commercial importance on this coast, since
pelagic fish are simply not caught at all. (No doubt plankton studies will be
of still greater importance if sardine and other pelagic fish ever become of
commercial significance, an achievement much to be desired.) Nevertheless
certain creatures such as the fish Apogonops anomalus, which is one of the chief
foods of the Tiger Flathead (Neoplatycephalus macrodon Ogilby, the most
important trawled fish of New South Wales), have been found with the stomach
contents entirely planktonic in nature. On certain occasions, the copepod
Calanus brevicornis was present to the exclusion of almost anything else, whilst
both adult and larval Nyctiphanes australis were abundantly present in the
stomach at other times when these species occurred in the plankton. But the
copepod C. brevicornis has never been common in our plankton catches, so that
either it occurs in shoals, and is deliberately selected by Apogonops, or it has
been missed by the net catches taken nearer land,

What information is at hand regarding the passage of shoals of pelagic fish
up our coast seems to indicate that these occur in the winter and spring months.
Calanus brevicornis was taken in Apogonops stomachs in October, whilst big
hauls of other large copepods have been taken in August, so that there is
already some evidence linking the occurrence of planktonic crustacea with the
movements of pelagic fish.

BY W. J. DAKIN AND A. COLEFAX. 215

The two most abundant types of fish egg which can be recognized in our
catches are: (1) an oval egg which is undoubtedly that of an anchovy, and (2) a
spherical egg of some clupeoid fish. The latter has been present in great numbers
in June and July—the young on hatching would just be ready for the phyto-
plankton maximum beginning in August. Apart from these two fish eggs, there
are several unrecognized species of pelagic eggs which are most common in
the summer catches (January to March). This fits in with the little that is
already known about the breeding season of some of the more common jnarine
fishes. It is unfortunate that in this respect so much stands to be done on the
Australian Coast.

Behind the fluctuations in the production of animal plankton, is the interesting
periodicity of the plants, and regarding this a little more may be added.

The onset of the diatom production in Arctic and Antarctic Seas is probably
to be correlated entirely with the occurrence of, and increase in, light after the
darkness of winter. In many temperate seas, and certainly in our own coastal
waters, the daylight is sufficiently strong for good diatom production at any
time throughout the year, even during the season of shortest days, and certain
diatoms are indeed always present. In this connection, we may also recall the
onset of reproduction of Gymnodinium in July, when the days are not far from
their shortest. Temperature has frequently been discussed in connection with
spring maxima, especially since the beginning of a spring maximum is often
coincident with the turn of the temperature “curve” from a downward grade to
an upward slope. It has been found, however, that this is not always the case,
and Marshall and Orr draw particular attention to the fact that the spring
increase may start before the temperature change, whilst one of the conspicuous
diatom species in the maximum studied by them in Scottish seas, flourished at
2°-3° C. on the Norwegian Coast, at 8°-9° C. in Loch Striven and occasionally is
found in the tropics.

Food substances are apparently present in abundance before the spring
increase of the diatoms commences. In some places the spring increase has
been correlated with melting snows and additions of fresh water and salts to
the sea. There is nothing of this sort in our region and nothing to show, up to
the present, that any chemical change takes place in the constitution of the
seawater which might lead to the onset of the spring increase. Marshall and Orr
in Scotland, and Atkins at Plymouth, after a consideration of the various possible
factors, are inclined to regard the intensity of the sunlight as the primary causal
factor. In regard to this, however, it may be pointed out that there is a big
difference between the length of the winter days and the summer days in the
regions where most plankton calendars have been worked out, and it is not
unusual for meteorological conditions to bring about still further poverty of
sunshine during the winter season when the days are shortest. Compared with
these conditions, any increase of daylight at Sydney when August is compared
with June seems but a small factor, for the duration of daylight only ranges
between 9 hours 53 minutes and 14 hours 25 minutes throughout the year.

Steuer affirms that the seasons of the great diatom maxima recede from the
warmest months of the year as one approaches the tropics. Bigelow confirms
this for the Atlantic Coast of the United States. If this is correct, and actually
our Sydney dates for the maxima are not in conflict with it, then latitude seems
an important factor in determining the onset of diatom reproduction, quite apart

216 MARINE PLANKTON OF THE COASTAL WATERS OF N. S. WALES, 1,

from the well defined oceanographical conditions associated with more extreme
limits of latitude—i.e., conditions peculiar to the equatorial as contrasted with
Arctic and Antarctic waters. And, so far as one can judge at present, this
latitude difference chiefly implies differences in light, and perhaps temperature,
although the latter is not so much a function of latitude as is intensity of light
and length of day, for ocean currents may be responsible for considerable
differences in sea temperature at the same latitudes. Unfortunately, “plankton
calendars” from tropical and subtropical regions are not sufficiently numerous and
the relation of latitude to the plankton rhythm requires further elucidation yet.

TABLE

The chief Diatoms of the N.S.W. plankton.

Jan. | Feb. | Mar. April. May. | June. July. Aug.
4 20 12 27 5 14 7 OW el Operate | 26M |econe tee les eats 2 8
= = = | = |
Asterionella japonica .. | — 6,600) 11,440 800 373,700 2,200 250} 400: x 4,000) 46,520
Bacteriastrum sp. a x 600 x
Biddulphia mobiliensis .. i 400 600} 6,600) 6,480
op jos | be ee
Coscinodiscus concinnus . . 5%
33 group ae x 6,600 3,500 600) 1,600 3,300} 1,600
Cerataulina sp. 55
Chaetoceras sociale We 440} 6,000
50 SDaa ie oe 880} 2,700) 12,320 19,800} 5,000) 48,180 2,000| 7,260) ~ | 1,500) 6,160) 10,080
Climacodium frauenfeldtii 440 600) 2,200; * 1,600) 500 700 230 s x
- biconcavum. . 250 °
Corethron hystrix : 6
Dactyliosolen tenuis ye x 200 200 & 300
Dityllium brightwelli : P 600} 7,480) 10,800
Eucampia zoodiacus a0 440 S
Guinardia flaccida ae 440 10,000
Hemiaulus hauckii se 400
Lauderia borealis ate ? 400
Leptocylindrus danicus .. 158,000! 22,000 i 55,500
Nitzschia seriata a 3,080 e ? 3,000} 880) 25,500/400
59 longissima A ; 700
Planktoniella sol .. ws 800} 220 900
Rhizosolenia alata .. |4,840 24,000 900} 200 300
re delicatula .. 440 :
on robusta Si 600
a sp. ae 17,400); 12,320) 22,440) 440 15,620; 1,900) 11,000,200 600; 200 1,200 720
* stolterfothii 2,700} 440 7,000 900 720 ©
Stephanopyxis sp. a 400
Streptothrix thamensis .. 400 400 700 400 900} 1,600, 3,600 ©
Thalassiosira condensata 2,000 4,500 3,600} 400 172,800 ©
3 rotula oo Wabsecsto) 1,200 750 300 300 13,680 |
ef subtilis ae x 600 x
3 sp... 32 6,600 400 | 22,000
Thalassiothriz nitzchioides 880 880 900)
Oscillatoria My Pe 800 ? 5,280) 900) 4,500 900

*Calcaravis.

I.

BY W. J. DAKIN AND A. COLEFAX. 217

As a matter of fact, there is much more to be investigated in the relation
between light and phytoplankton. More accurate estimations of the intensity of
light at depths in the sea are now being obtained as the result of the use of
“light cells’ and direct photometric methods. From these it appears that only
6-62% of the sunlight may reach a depth of 20 metres (it may, of course, be
much less) and only 0:7% a depth of 40 metres. The spring diatom maximum in
high northern latitudes commences when the light cannot be very intense below
the surface. The amount of light which is optimal is another unknown, and the
matter is probably more complicated than has been recognized. It has been

Seasonal Range 1931.

Aug.

112,700] 57,860
8,406, 440
5,600/ 800
8,400} 900
17,500] 8,800
3,520
12,600, 900
700
450
700| 1,320
500
3,500
9,000
900
1,400| 400
8,400 | 44,660
6,300| x
DG
x

Sept. | Oct. Nov. Dec. Jan.
5 12 20 3 17 25 1 13 15 BA WAS!) 26 2 8
x | 22,630] 23,100) x 6,600,
700 200 |
x | 1,460) 1,960) 880! 4,500 10,000
1,600 1,600
2,200 8380 4,000
800 3,600, 3,000
450) 304,000)
47,450, 64,020 |5,720 |365,560| 16,500) 21,120) x 6,600 650) 11,000 19,000
é : 440 x
3
$ 4,400 400
3 730 2,000} 2,400
+ | 12,410; 6,600 440
A | 2.800) 4,800 6,700} 800; 3,000 300) 600) 113,520 450
2 4s
2 450 3 1,200, 400
cs ? 100 es
3 1,400] x 2,200} 5,000 3 12,000 78,320 1,200
= | 3,500, 3,500) 600 a 600 3,200 1,200
5
S
700 9,240 2 8,800 19,000
880 =
8 | 17,000*
3,500} 1,300, 450 200) 26,400 120,000] 28,000} 4,400 3 3,200 15,840
600) 2,400 rae 800,
43,800} 50,000 2,200) 2,500 4,400 1,500 8 z
ae 440
2,100 880 400 ong 400
87,600, 222,000, 7,040 4,000 £8 1,200 800
2
4,200 sp. ? I
2.200| x 13,000 “
= 3,200 1,200
o

YWith parasite Richclia.

Feb.
|
28 10
650
|
2,000 200
| 85,800
440
220) 220
} 220
|
| 2,200
| 5,600
220
1,000+
850
o
= 5 440
>)
238
Os
~
24,000 15,600

218 MARINE PLANKTON OF THE COASTAL WATERS OF N. S. WALES, i,

TABLE

Q The seasonal distribution of the chief zooplanktonic forms in
; The figures are comparative only and indicate the number of organisms counted in

each date. It is possible also to compare the numbers of different species of

J and larvae are enumerated from
1931.
| Jan. | Feb. | Mar. | Apr. | May. June. July. Aug.
4 20 12 PAC 5) 29 7 10 1 26 21 2 18 2
PROTOZOA
Favella campanula a0 600 440 250
Codonellopsis ostenfeldti 440 250 200
Tintinnopsis radiz i. j 1,320 220 200 | 1,200
Gymnodinium sp. ae | 1,600
Peridinium sp. .. .. | 4,400} 1,200] 3,080 1,980] 4,000} 2,400) 600; 880] 4,200
Ceratium tripos oe 880} 1,800 440 1,400 200} 730} 1,200 250 880
as furca .. .- | 1,760) 33,300 880 600 200) 5,000 600 440
a ipeertuscayecy we tae 880 2,200 700
Dinophysis tripos ot: 600 200 | 220 1,600 300
COPEPODA |
Acrocalanus gracilis .. 700 1 x x x 140 x |
Acartia clausi .. .. | 11,480 560] 5,740] 2,240) 280} 2,380,1,820] x 140) 700 280 2,380) 700} 700
Calanus pauper. . ae 840 700 x x x
Candace pectinata aie xox
ty ethiopica mee x< x x
Eucalanus sp. immature x x 140 |
Euterpe acutifrons a 140 560 x 420 x x 700; 140} 420
Labidocera acuta ae x x x x x x x x xe x
sy cervi ate oe x x x<
Mecynocera clausii
Microsetella rosea as 560} x x 1,400) 560; 560 280) 140) Naup.
Oithona setigera. . |
py oculata eee x
=f plumifera ee x 560} 280 140) x
on tenuis aN 420 sp. 840) 1,400] 560
Oncaea venusta os 140; x x 280 x 560} 140 700| 420) 280) 140 140
Onychocorycaeus catus . . 3,780| 840) 140 1,680 1,120} 420) x 140; x x
Paracalanus parvus .. 700 980) 5,600) 840) 7,140 4 4,340} 420) 17,500) 1,400} 1,960) 11,200 700; 280
Temora turbinata fi 980} 3,080) 15,540) 1,820} x x 140} 1,960 700 420 420 140
OTHER CRUSTACEA |
Evadne nordmanni
» spinifera .. | 6,720] 1,400] 4,060] 1,120 140 280| x
¥e tergestina a 280] 3,500) 4,060} 1,260 140} 2,660) 2,940 1,680! 620 140
Podon polyphemoides .. 560
Penilia schmachkeri .. x 1,260 420} 140 140) 1,120) 140 280} 2,520 140 |
PTEROPODA
Creseis virgula ..
UROCHORDATA
Oikopleura sp. .. aie x x 700; 1,560) 560 560} x x 280 140; x 560} 140
Fritillaria pellucida .. x 140} 2,240 140 2,260 140] x
Thalia democratica An WHO WECo UIEVCS AVEC v.c. v.c. V.c.
EG@s AND LARVAE
Anchovy eggs fe 140 }
Other fish eggs ats x x x 280] x 140
Penaeid eggs .. ive : 560} 1,120 2,240} 2,380 1,400) 420 560
Lamellibranch larvae .. 1,120 x
Veliger larvae 60 jled) 36 1,400 980 980 140 140 140] x
Ophiopleuteus .. “a 2,380 14,140} 140
Nauplii—barnacle ar 280 x< 140 140] 560
Zooea .. oie ae x x x x 280

Ti.

BY

Ww.

J. DAKIN

AND

New South Wales waters off Port Jackson during 1931.
catches made, so far as any particular species is concerned, in exactly the same way on
Protozoa, Copepoda, other Crustacea and Urochordata on any date, but the eggs
catches made with a different net.

A. COLEFAX.

219

1931. 1932.
rey
Aug. = Oct. Nov | Dec Jan Feb.
w
LZ 20 20 3 17 25 | 1 15 22 a 23) 26 2 8 28 10 23
| Night.
800
400
700
700; xX 3,200
2,800} x x 4,050 | 3,500) 1,500; 67,500} 1,800} 30,000 | 3,500 x 2,500} 3,000 450| 5,400
x x x 2,450 880 880 400 400
440 3,000 |1,000! 80,000) 1,760} 3,600 | 10,800 600; 3,500 400 4,000
450] xX x 1,000} x
450} 2,000; 900) 14,080 600} 1,200 220
140 x
560) 14,200} x 2,940} 10,000} 140) 18,000) 900) 6,500 840} 2,400) 2,100} 8,000} 6,580 980} 4,201) 920
2 x 140
140 x
140 x x
>< 3,080 140 280 280 140
x
x 140 140 *
280 x
x sp. 1,120 x
? 420 280
560 140 x 2,000; 140 280 280 720 420 980| x x 1,400
140} x 140 420 140 280
x x x 140 x x
x |384,440) x 420} 3,500) 1,000) 32,100) 4,000) 10,000 420) 5,000| 8,260| 17,000) 18,500) 28,840} 17,000) 840
x 7,140] x 1120) 280 280| 280 140 280 840 140
140 140 420 280 8,000) 3,400 840 140} 4,060 280
280 980} 1,960} 980 560! 11,200} 1,680} 8,200) x
280) 1,820 x x 500 3,080} 8,000} 5,000) 7,400 700 140) 3,000
x 6,020; 9,000) 2,800 280
x 140 x x 700} 1,260 420 140 2,800
140 36
1,640 560 280 560| xX 560} 140; 1,260 420 140 280 140 980} 2,160| 1,680
840 980 Deep X x 140 140 420) 600
40,000
280 140; o.c x
few < x 60 X rare | Vv. rare x
x x x x x Xe 140} x x 420
x 280
280 140 560
840
140 4,010 140} x 140 1,400
140 140 840 140 x
2,240 800 x 700 600 2,000; 980 140
x 140) x

220 MARINE PLANKTON OF THE COASTAL WATERS OF N. S. WALES, i,

shown for example for the higher plants (by Popp, Amer. Journ. Botany, Vol. 13,
1926) that the blue-violet end of the spectrum is necessary for normal vigorous
growth (ultra-violet rays are not indispensable) and yet photosynthesis proceeded
at a more rapid rate at the red end of the spectrum. Illumination with the full
spectrum was more important than light intensity with certain wave lengths
removed. Now, so far as we are aware, nothing of any consequence has been
determined regarding the effect of different wave lengths on the growth of phyto-
plankton, yet the sea is a medium which differentially filters the longer wave
lengths compared with shorter. Moreover, it is just in regard to the ultra-violet
and blue-violet rays that there is greatest change in illumination between winter
and summer in temperate latitudes.

These facts have been emphasized because it is difficult to escape the conclusion
that there is more in the control of plankton maxima than immediate environ-
mental factors. During the plankton maxima in the English Channel the phosphate
and nitrate content of the sea diminishes until the phosphate becomes practically
zero in the early summer and the nitrate almost as low. It has been suggested
that it is this reduction in nutrient salts which brings the diatom maximum to
an end. Possibly this may be important in regions where essential nutrient
salts are reduced below the minimum by enormous plant reproduction. It is
curious, however, that the diatom curves should be so similar in such diverse
places. Is the seasonal distribution of nitrates and phosphates always the same?
There are other difficulties, too, in applying these theories. Marshall and Orr
showed that the diatom increase came to an end before the phosphate was all
utilized. And how is the autumn peak to be explained, or the luxuriant develop-
ment of dinoflagellates and blue green algae immediately after a diatom maximum?

In our case the analyses do not show a big reduction in phosphate, or
nitrate, after the spring maximum, and one is impressed all the more by the
rhythmically occurring reproductive periods of different species. This periodicity
must be the result of the interplay of internal factors and environmental factors,
and it is even conceivable that the date of onset of a maximum may be deter-
mined, not only by the occurrence of favourable environmental factors, but by
the previous year’s history of events. It may be worth while in this connection
to make a comparison with the yearly periodicity of land plants (although once
again there seems little enough experiment to elucidate this subject). Certain
of the temperate plants exhibit seasonal periodicity after years of growth in
tropical or subtropical areas notwithstanding an environment which is practically
uniform. It is true that, in the case of the diatoms and other phytoplanktonic
forms, we are dealing with single-celled organisms with quite a short life, but
it is not unusual by any means for spores, eggs, etc., to require a certain resting
period before they are capable of further development. In any case, we must
postulate the existence of a delicate adjustment between the organism and its
environment exhibiting itself in the control of growth and reproduction. In no
other way, for example, can we account for the remarkable burst of reproduction
of the Gymnodinium which colours Port Jackson water red for two or three
weeks every year and which times its occurrence with close adherence to the
calendar. In this connection, too, it is significant that the maximum formed by
the chief species of Rhizosolenia in New South Wales waters bears a time
relation to the onset of the spring diatom maximum (the rise in numbers of

BY W. J. DAKIN AND A. COLEIFAX. 221

Thalassiosira, Chaetoceras and Asterionella) similar to that observed in British
waters. This is particularly interesting in view of the fact that the seasons are
alternate (Southern Hemisphere) and that the sea temperatures at our station
are at all times higher than those of British Seas. We are inclined, therefore,
to regard the periodicity of the plankton as controlled no less by internal factors
than by the directly potent factors of the environment in the fullest sense. The
external factors may be of greater importance than the internal in “touching the
trigger’ for the vernal maximum, for example, but the succession of different
species, and the whole aspect of the rhythmic cycle of change is probably due to
the reaction of one organism on another, and to inherent characters which are
little understood.

The first long period study of the plankton conditions in Australian Seas
has, up to date, provided material of considerable interest from the point of view
of geographical distribution, and it has indicated the occurrence of a regular and
definite seasonal change in the plankton of our seas—a change which is both
qualitative and quantitative. Since many commercial fishes feed upon plankton
during at least some part of their life, there is reason to believe that migratory
movements of fishes will be found to be related to these seasonal planktonic
conditions in our southern Australian waters. It is necessary, however, to
emphasize distinctly the limitations and difficulties of this present work. No
complete knowledge of either the hydrographic or the biological conditions of
the coastal waters of the Tasman Sea can possibly be obtained without the
organization of such work on a much bigger scale and over a large area. The
continuation of our work should provide the foundation for such a scheme.

Literature Consulted.
ALLEN, W. E., 1927.—Surface Catches of Marine Diatoms and Dinoflagellates from
Pacific High Seas in 1925 and 1926. Bull. Scripps Inst. Oceanography, Tech. Series, i,
No. 12, 1927.
, 1928a.— Quantitative Studies on Marine Diatoms, ete. TIbid., Tech. Ser., i,
No. 15, 1928.
, 1928b.—Review of Five Years of Studies of Phytoplankton at South California
icra ods, Lech) Ser 1, Now 26s) 119128)
Bravby, G. S., 1883.—Copepoda. Challenger Reports, Vol. 8, Pt. 23.
, 1918.—Copepoda. Aust. Antarctic Exped., Series C,. Vol. v, pt. 3.
VAN BREEMEN, 1908.—Copepoda. Nordisches Plankton, viii, 1908.
Cuiaus, C., 1877.—Kenntniss des Baues der Polyphemiden. Denk. Akad. Wissenschaften,
Bd. 37, Wien.
Cueve, P. T., 1901.—The Plankton of the North Sea and Skagerak in 1900. Kung.
Svensk. Vetenskaps-Akad., Handlingar.
Dawu, Maria, 1912.—Die Corycaeinen. Pt. i of Die Copepoden der Plankton Expedition,
Vol. 2.
ESSENBERG, C. E., 1926.—Copelata from the San Diego Region. Univ. of Calif. Publica-
tions.in Zoology, Vol. 28, No. 22.
ESTERLEY, C. O., 1905.—Pelagic Copepoda of the San Diego Region. Univ. Calif. Pub. in
Zoology, Vol. 2, 1905.
, 1906.— Additions to the Copepod Fauna of the San Diego Region. TIbid., Vol. 3.
, 1911.—Third Report on the Copepoda of the San Diego Region. JIbid., Vol. 6.
FARRAN, G. P., 1910.—Copepoda. Brit. Antarctic Exped. (‘‘Terra Nova’). Zoology,
Vol. 8, No. 3.
Fisu, C. J., 1925.—The Seasonal Variation of Plankton of the Woods Hole Region.
Bull. U.S. Bureau of Fisheries, Vol. 41, 1925.
GIESBRECHT, W., 1892.—Die Pelagischen Copepoden. Flora und Fauna des Golfes von
Neapel, Monograph 19.
GRAN, H. H., 1905.—Diatomeen. Nordisches Plankton, 19. Kiel, 1905.
, and ANGsT, E. C., 1931.—Plankton diatoms of Puget Sound. Public. Puget
Sound Biolog. Station, Univ. of Washington, Vol. 7, 1981.
iat

222 MARINE PLANKTON OF THE COASTAL WATERS OF N. S. WALES, i.

HANSEN, H. J., 1899.—Die Cladoceren und Cirripedien der Plankton Expedition. EHrgeb.
der Humboldstiftung Plankton Haped., Bd. 2. Kiel, 1899.

HERDMAN, W. A., 1918.—Spolia Runiana. Distribution of Certain Diatoms and Copepoda
throughout the Year. Jowrn. Linn. Soc., Vol. 34, 1918.

KARSTEN, G., 1905.—Das Indische Phytoplankton. Wiss. Ergeb. der Deutschen Tiefsee
Eaped. (‘Valdivia’), ii. Jena, 1905.

KoKUuBO, S., 1932.—Records of Oceanographical Waters in Japan. Nat. Research Council
Japan, Vol. 4, Tokyo, 1932.

KrAmer, A., 1894.—On the most frequently occurring Pelagic Copepoda and Cladocera
from the Hauraki Gulf. Trans. N.Z. Inst., Vol. 27.

Koroip, C. A., and CAMPBELL, A. S., 1929.—Conspectus of Marine and Fresh Water
Ciliates belonging to the Sub-order Tintinnoina. Univ. Cal. Pub. Zool., Vol. 34,
1929.

Koro, C. A., and TAGE SKAGSBERG, 1928.—Dinoflagellata. The Dinophysoidae. Mem.
Mus. Comp. Zool. Harvard, Vol. 51.

,and Swezy, O., 1921.—Free Living Unarmed Dinoflagellates. Mem. Univ.
California, Vol. 5, 1921.

Lesour, M., 1930.—The Planktonic Diatoms of Northern Seas. Ray Society Monograph,
No. 116, London.

LEMMERMAN, 1903.—Flagellata and Silicoflagellata. Nordisches Plankton, 21, Kiel, 1903.

LOHMANN, H., 1899.—Appendicularien. Hrgeb. der Plankton Expedition der Humboldstif-
tung, Bd. 2, Kiel, 1899.

MARSHALL, S. M., and Orr, A. P., 1927.—The Relation of the Plankton to Some Chemical
and Physical Factors in the Clyde Sea Area. Journ. Marine Biol.. Assoc., N.S. 14,
No. 4.

Marcatr, MAyNArD M., 1918.—The Salpidae: A Taxonomic Study. Smithsonian Institu-
tion, U.S. Nat. Mus. Bull 100.

MUuuer, G. W., 1906.—Ostracoda. Ergebiisse der Deutsch. Tiefsee Exped. (“Valdivia’’),
Bd. 80, Jena, 1906.

Murray, Sir J., 1895.—Summary of Results. ‘‘Challenger’’ Reports, Pt. i, 1895.

PAULSEN, O., 1906.—Peridiniales. Nordisches Plankton, xviii, Kiel, 1906.

PELSENEER, P., 1888.—The Pteropoda. ‘Challenger’? Report, xxiii, 1888.

ROSENDORN, Iusg, 1917.—Die Gattung Oithona. Wiss. Ergeb. der Deutschen Tiefsee
Eaped. (“Valdivia’), Pt. i.

SCHIEMENZ, P., 1906.—Die Pteropoden der Plankton Expedition. Leipzig, 1906.

ScuHtrr, 1895.—Peridineen der Plankton Expedition. Hrgeb. der Plank. Eaped. der
Humoboldstifitung, Kiel, 1895.

Scorr, THos., 1894.—Report on the Entomostraca from the Gulf of Guinea. Trans.
Linn. Soc., Zool. Vol. vi, Pt. i.

Scorr, ANDREW, 1909.—The Copepoda of the Siboga Expedition. Pt. i. Siboga Expéditie
Monograph xxixa.

SEYMOUR-SEWELL, R. B., 1912.—Notes on the Surface-living Copepoda of the Bay of
Bengal. Rec. Ind. Mus., Vol. vii.

Sm1TH, Ep. A., 1888.—Report on the Heteropoda. ‘‘Challenger’’ Report, xxiii, 1888.

STEUER, A., 1911.—Leitf. der Planktonkunde. B. G. Teubner, Leipzig, 1911.

TescH, J. J., 1906.—Die Heteropoda der Siboga Expedition. Siboga Eapéditie Mono-
graph li, Leiden, 1906.

THOMPSON, Isaac, 1890.—Copepoda of Madeira and the Canary Isles, with Descriptions
of New Genera and Species. Journ. Linn. Soc., Zool. Vol. xx.

THOMPSON, Issac, and Scotr, ANDREW, 1903.—Copepoda. Ceylon Pearl Oyster Fisheries
Report, Part i.

WHITHLEGGE, THOS., 1891.—On the Recent Discoloration of the Waters of Port Jackson.
Rec. Aust. Mus., i, No. 9, 1891.

EXPLANATION OF PLATE VII.
1.—A Ctenophore, n. sp., from the New South Wales coastal plankton.

2, 3.—Some common fish eggs. (2) Anchovy eggs. (3) Clupeid eggs, species
doubtful. ;

4.—Thalia democratica.

Proc. Linn. Soc. N.S.W., 1933. PLATE VIL.

1.—A new species of Ctenophore. 2.—Anchovy eggs. 3.—Clupeid eggs.
4.--Thalia democratica.

co

NOTES ON NEW SOUTH WALES AND QUEENSLAND ORCHIDS.
By the Rrv. H. M. R. Rupp, B.A.

(Two Text-figures.)

[Read 28th June, 1933.]

A. Relations between certain forms of Dendrobium.

For some years past botanists and orchid-fanciers alike have been puzzled
over the relations between Dendrobium speciosum Sm., D. gracilicaule F.v.M.,
D. Kingianum Bidw., and certain allied forms which vary considerably, at times
being sufficiently distinct to give the impression of independent species, and at
times apparently justifying the opinion that they are merely intermediates due
to hybridization between the above-mentioned species. These forms are at present
known as D. delicatum Bail., D. Kestevenii Rupp, D. speciosum var. nitidum Bail.,
and D. speciosum var. gracilimum Rupp. It seemed worth while to try and clear
up this confusion and, though I can scarcely claim to have done this, the results
of my examination of a Jarge number of specimens from various sources may at
least serve to simplify the problem for all who are studying it. The difficulties
have been intensified by the fact that the late F. M. Bailey apparently left no
herbarium types of D. delicatum and D. speciosum var. nitidum, and consequently
these names have been bestowed upon different forms without justification: e.g.,
white-flowering forms of D. Kingianwm have passed for D. delicatum. In 1931,
Mr. C. T. White, the Queensland Government Botanist, sent me the Brisbane
Herbarium specimens labelled D. speciosum var. nitidum, and with one exception
I found the flowering specimens identical with a form to which several years
ago I gave the name var. gracillimum. The exception came from Tambourine
Mountain, and in September, 1932, Mrs. H. Curtis sent me abundant living material
from that locality. I found this form to be quite distinct from var. gracillimum,
and it appears to me to conform in every respect to Bailey’s description of var.
nitidum. In my opinion this should rank as a species. The stem is still more
slender than that of var. gracillimum, but the flowers are much larger, and pure
white except for faint markings on the labellum. The latter is of very thin
texture, quite distinct in form (Text-fig. A, 8) from that of D. speciosum
(Text-fig. A, 1, 2), and the sinus between each lateral lobe and the mid-lobe is so
deep that the latter easily breaks off in handling unless one is careful.

D. speciosum var. gracillimum was described and discussed in These
PROCEEDINGS, liv, pt. 5, 1929. I agree with those who feel the great dissimilarity
in appearance between this and other forms of D. speciosum to be a stumbling-
block in the way of accepting it as a mere variety. I pointed out that the flowers,
however, in the typical form of the variety, are indistinguishable from those of
some of the small-flowered robust D. speciosum, and this fact, which is corroborated
by other workers, seems to debar it from specific rank. It has been conjectured
that var. gracillimum criginated by hybridization between D. speciosum and

224 NEW SOUTH WALES AND QUEENSLAND ORCHIDS,

D. gracilicaule. The general appearance of the plant favours this theory, and
Mr. A. G. Hamilton, of Chatswood, has a plant with heavily-spotted or biotched
flowers—even the labellum (Text-fig. A, 5). Mr. Hamilton’s flowers, however, are
exceptional, and the form of the labellum is very peculiar. It will be seen (Text-
fig. A, 4-7) that the labellum of this orchid is subject to considerable variation:
No. 7 belongs to a flower of which specimens were sent independently by Messrs.
E. Slater of Bullahdelah, and R. Leaney of Chatswood, under the name ‘‘cream-
flowered D. Kestevenii’. It is, however, quite unlike the D. Kestevenii labellum,
and the whole flower agrees with gracillimum, though the stems are shorter and
somewhat sturdier than the type.

D. delicatum Bail. remains, for practical purposes, an unknown quantity.
Mr. F. A. Weinthal, of Chatswood, sent me a plant obtained in Southern Queens-
land some years ago, which he believes to be Bailey’s species. The labellum is
shown in Text-figure A, 12. No. 11 is from a plant in the bush-house of Mrs.
C. A. Messmer, Lindfield. Mr. Weinthal’s plant is of a more robust type: the
flowers are not unlike, but the two labella are very different. No. 10 is from
Mrs. G. Annand, Lismore, labelled ‘‘white Kingianum’’, but the plant seems to me
too robust and hard of texture for that species. No. 9 (Mrs. Messmer) is a typical
D. Kingianum, but the species has many varieties. It can scarcely be doubted
that 10 and 11 show close affinities with 9: this is by no means so obvious with
12, which seems nearer to the series following. Nos. 13 to 18 are all from

18

Text-fig. A.—lLabella of certain forms of Dendrobium in New South Wales

and Queensland. 1. D. speciosum Sm., a small-flowered form; 2. D. speciosum,

a large-flowered form; 8. D. gracilicaule F.v.M.; 4, 5, 6, 7. D. speciosum var.

gracillimum Rupp; 8. D. speciosum var. nitidum Bail.; 9. D. Kingianwm Bidw. ;

10. D. Kingianum ?; 11. D. delicatum Bail. ?; 12. D. delicatum ?; 13 to 18.
D. Kestevenii Rupp.

plants supposed to be D. Kestevenii. We may call 13 the type, since it is from
the plant originally sent to me from Bullahdelah by Dr. H. L. Kesteven, after
whom I named it. Nos. 14, 15, and 16 are from Mr. F. Fieldsend, of Hast Maitland:
17 is from Mr. E. Slater, and 18 from Mr. R. Leaney. All the plants originally
came from Bullahdelah. Even if No. 12 should prove to be the genuine D. delicatum
of Bailey, it is not quite identical with any of those known as D. Kestevenii: and
the stems or pseudobulbs are smoother and more slender. But the affinities are
so close and so obvious that my present view of the problem is, that it will be
found advisable to unite these two to constitute a single species. They cannot
justly be included under either D. speciosum or D. Kingianum, but they do appear
to come between those species. Further study, however, is desirable before such

BY H. M. R. RUPP. 225

a step is taken. I am frankly puzzled by Nos. 10 and 11, but in view of the
unmistakable “Kingianum influence’ in the labella, I should class them in that
species until we have further working data.

Note on Teat-figure A.—It will be seen that the labella differ so much in detail
that they are not safe guides even to a particular species. It is of interest to
observe the varying characteristics of the median line, which may prove of some
value in tracing affinities. Thus if we take 1, 3, and 4, we find a general resemb-
lance in outline, and the forms of the median line are identical. No. 12 is very
peculiar: the median line ends in three teeth, and has two triangular wings on
each side. In 13 it has one such wing on each side, and ends in two curious flaps.
In 10 it ends in two teeth. No. 2 is the labellum of a very beautiful D. speciosum
sent by Mr. E. Slater, with large flowers: the prominent markings on the labellum
are maroon. All the drawings are semi-diagrammatic, made from labella flattened
out.

B. Cryptanthemis Slateri Rupp.

Reference to the description of this genus and species in These ProcEEDINGS,
lvii, Parts 1-2, 1932, will remind readers that the original specimens were found
late in November, when all the flowers were more or less withered and shrivelled.
Though it was possible to soften a few of them sufficiently to identify the different
parts, I suggested that the description then given would probably require to be
supplemented when fresh specimens, at a less advanced stage of development,
should be discovered. I had hoped to visit Bullahdelah, the scene of the original
discovery, in the spring of 1932, with a view to examining plants in situ, but was
unable to do so. Early in October, however, Mr. Slater sent me further specimens
from Bullahdelah. Unfortunately they were somewhat damaged en route, and
as I was away from home when they arrived, they were immersed in water until
my return; this freshened them up, but did some injury to the minute and
delicate details of the flowers. In view of the fact that these specimens were
six weeks earlier than those of 1931, it was disappointing to find that of the four
capitula sent, three had only flowers well past maturity. The fourth, however—
a slender one with few flowers—was in good condition and fhe flowers had not
withered. It was at once evident that the figures accompanying the criginal
description, drawn from withered flowers “restored” as accurately as circumstances
permitted, did not adequately represent living flowers.

The living flower of Cryptanthemis, in fact, bears far more resemblance to
that of the Western Australian Rhizanthella than is apparent in withered
specimens. In the latter the segments have the appearance of being membranous
and flaccid, inclined to diverge from one another. In the living state the whole
flower is very succulent, and the segments are thick and set very closely together
in an erect position. The paired sepals in particular are rigidly erect, with their
narrow and prolonged anterior portions inclined at an angle towards the centre
of the capitulum. These anterior portions, though relatively slender, can scarcely
be termed ‘“filiform’’. At their bases the paired sepals are gibbous, but as the
flower withers and the ovary enlarges, this feature is lost. Reference to figure 5
in the original description will show the ‘median line” of these sepals to be not
truly median, but slightly to one side. This is correct, though in the figure cited
the sepals are shown with their apices downwards, which is never the case in
the living flower. Text-figure B, 8 shows the real character of this “median”
line, which forms a ridge along the exterior convex surface of the sepal. If a

226 NEW SOUTH WALES AND QUEENSLAND ORCHIDS,

cross-section of the latter be taken, it has the form. of two sides of a scalene
triangle, the ridge being the apex.

Viewed from the front of the flower, these rigidly erect paired sepals, concave
within, form with the dorsal sepal a kind of box protecting the petals, labellum,
and column. The margins of the petals are very irregularly denticulate. In the

Text-fig. B.—Outline sketches of Cryptanthemis Slateri, Oct., 1932. 1. A plant

with three branches, the main rhizome having a capitulum of living flowers;

2. flower from front; 3. flower from the side; 4. flower from the front with

sepals removed; 5. labellum and column from the side; 6. effect of pushing

labellum away from column; 7. column from the front; 8. dorsal view of one
of the paired senals; 9. a petal (2 to 9 variously enlarged).

text-figures of the original description the segments and other parts of the
individual flower are shaded dark; but the living flower (except for a brownish
patch on the back of the column which is not a constant feature) is almost wholly
white, nor could I see any tendency to discoloration after exposure to light,
such as Dr. Rogers records of Rhizanthella.

The labellum is larger than it appeared to be in the 1931 withered flowers,
but is still relatively smaller than that of Rhizanthella. Its whole surface is
glandular-rough, and I think (it is difficult to be sure) that the apparent minute
denticulation of the margins is really due to this glandular roughness of the
surface. The claw or stalklet attaching the labellum to the column-foot, in all
the living flowers examined, appeared to me to have lost its power to function—
probably by the prolonged immersion in water related above. In each case the
labellum was erect with its ventral surface against the front of the column
(Text-fig. B, 5), but directly it was pushed away it fell down on the floor of its
“box” (Text-fig. B, 6), the claw being apparently unable to support it. In
undamaged flowers it seems probable that the labellum is normally held in a
horizontal attitude with its apex projecting between the paired sepals.

BY H. M. R. RUPP. 227

The column is short and stout, with a relatively large stigmatie plate.
Examination of its details in these water-soaked specimens was difficult, especially
as they reached me during a temporary sojourn on the inland plains of New
South Wales, where I had few facilities for such work. As far as J could judge,
the two columnar appendages mentioned in the original description are analogous
to the “horns” on the front corners of the column-wings in some species of
Pterostylis: they appear to rise from the rostellum, and are straight, or only
very slightly curved. In the outer flowers of the capitulum the anthers had
disappeared, and there were traces of pollinia on the stigmatic plates.

Much yet remains to be learnt about this remarkable plant, and it is most
desirable that search should be made for it wherever Dipodiuwm punctatum, the
“associate” of Cryptanthemis, is known to occur. In this paper I have confined
myself chiefly to correcting and amplifying the original description as far as is
possible at present. I may add here, that notwithstanding the compact and
almost tubular appearance of the living flower, and the “dovetailing” of the sepals
to form a sheltered chamber for the inner parts, there can be no doubt that the
sepals and petals are free.

There appears to be remarkable variation in the form of the plant itself.
Of the nine specimens received since the first discovery, three were less than
two inches long, but thick and compact. The others were elongated and slender.
One of the 1932 specimens had three branches, the lower portions of these being
without bracts.

C. New Records of New South Wales Orchids.

1. Caleana Nublingii Nicholls—Described by Mr. W. H. Nicholls (Vict. Nat.,
May, 1931), but not hitherto recorded in any New South Wales publication.
Discovered by Mr. HE. Nubling at Bell, in the Blue Mountains, 27th December,
1930. Near C. minor R.Br., but differs in its blunt, pear-shaped labellum and other
details.

2. Caladenia alpina Rogers.—Specimens in the National Herbarium, Sydney,
collected by the late Mr. R. H. Cambage at Queanbeyan, and labelled C. carnea,
undoubtedly belong to this species. Previously only recorded for New South
Wales by Mr. G. V. Scammell at Kosciusko.

3. Corysanthes unguiculata R.Br—Brunswick Heads, Aug., 1932, Mr. F.
Fordham. A very interesting record, extending the known range of this species
northward by 350 miles.

4. Thelymitra aristata Lindl.—Brunswick Heads, Sept., 1932, Mr. F. Fordham.
Growing among dense masses of Dendrobium Kingianum. Mr. Fordham supposed
it to be 7. longifolia, but though even smaller than the Keilor (Vic.) TZ. aristata,
it seems to me undoubtedly that species. The column-hood is yellow in front,
dark behind, with a broad V-notch; the buds expand readily, and the perfume is
strong. It is surprising to find 7. aristata associated with a Dendrobium on our
far North Coast. I was rather struck by the superficial resemblance of the
living specimen to D. Kingianum itself. The slender stem was curved; the colour
and perfume of the flowers were not unlike, and the dimensions about equal.

5. Lyperanthus ellipticus R.Br.—Peat’s Ridge, Mangrove Mountain, early
1932, Mr. H. Chapman. I do not know if this species has previously been recorded
on the northern side of the Hawkesbury River, but it is generally supposed not
to extend in that direction, and a definite record is therefore of value.

228 NEW SOUTH WALES AND QUEENSLAND ORCHIDS.

6. Diuris palachila Rogers.—Molong District, Sept., 1932 (W. H. Blakeley).
Mr. Blakeley’s specimens agree well with the South Australian type form.

7. Pterostylis Baptistii Fitzg—Mrs. C. A. Messmer has found this species at
Lake Tabourie, south of Milton, thus extending its range well to the south of
Jervis Bay. She has also recorded there Caladenia carnea R.Br. var. gigantea
Rogers; but though this is the most southerly record for New South Wales, the
variety has been identified (1932) at Airey’s Inlet, on the Victorian coast between
Port Phillip and Cape Otway.

Varietal Descriptions.

In order to comply with the international rules of nomenclature, the following
Latin descriptions of named varieties described in These Procerpines are
supplied:

DENDROBIUM SPECIOSUM Var. GRACILLIMUM (Vol. liv, part 5, 1929).—Scapi
gracillimi, 22-40 cm.-alti. Folia 8-16 cm. longa, non rigida, multo curva. Racemi
10-25 cm. Flores parvi cum segmentis brevibus.

PTEROSTYLIS OPHIOGLOSSA Var. COLLINA (Vol. liv, part 5, 1929).—Flos parvus,
pars superior fuscoruber. Galea breve acuta. Sepala lateralia brevia. Labellum
multo curvum.

PTEROSTYLIS ACUMINATA Var. INGENS (Vol. liii, part 5, 1928).—Planta robusta.
Flos quam forma typica semper multo major.

PTEROSTYLIS PUSILLA Var. PROMINENS (Vol. lvi, part 2, 1931)—Magnopere a
forma typica differt. Planta 10-30 cm. alta. Flores nutantes, saepe numerosi,
fuscorubri, a scapo prominentes.

CALADENIA DILATATA var. CONCINNA (Vol. liii, part 5, 1928).—Planta parva.
Floris segmenta omnia brevia, prope aequalia, acuminata. Labelli apex brevis-
simus, margines posteriores dentati.

THE SURFACE HISTORY OF MONARO, N.S.W.
By Frank A. Crart, B.Sc., Linnean Macleay Fellow of the Society in Geography.
(Plate ix; four Text-figures.)
[Read 26th July, 1933.]

In this paper, the normal landscapes found in part of the south-eastern
highlands of Australia are described, and regional surfaces of erosion are correlated
with those already determined in the Shoalhaven Valley. The plateau has a
terraced appearance which is held to be chiefly due to normal erosion in stages,
as opposed to the idea of differential uplift of a common surface: the stream
patterns have changed very little over long periods of time, and were evolved
from longitudinal streams modified, perhaps, by ancient depressions in the primary
surface. The northerly section of the Main Divide to the north of Monaro is
not of much tectonic consequence, but the Victorian main divide has a general
easterly extension towards the Pacific coast, and is convex to the ocean. In
addition, the Tertiary drifts of Kiandra are held to be an expression of changing
hydrography rather than of tectonic interference.

Introduction.

The name “Monaro” is sometimes applied to the whole extent of the higher
plateau in south-eastern New South Wales, but it belongs specifically to the
watershed of the Snowy and Murrumbidgee Rivers between Rule’s Point and
Nimmitabel. The more restricted sense of the word is used here, and most
attention is paid to a strip of country extending some 20 miles on either side
of the divide. The features of the Cooma district and the long northerly course
of the Murrumbidgee are being treated separately by Dr. W. R. Browne, to whom
due acknowledgment is made for references to his area, and for his exposition
of its features and problems. Former accounts include the admirable narrative
of Rev. W. B. Clarke (1860), the systematic work of Sussmilch (1909) and Griffith
Taylor (1910), and references by David (1908), Andrews (1910) and Browne
(1928), so the broad outlines of the surface features are already known, although
they have been studied with an eye to tectonic history rather than surface
classification, an aspect which repays detailed study.

Scenery and Topography.

The scenery and stream patterns fall into a few simple types with occasional
overlapping when the major controls of climate, rock character and elevation
combine in almost equal vroportions: thus the appearance of alternate basins
and narrows which is characteristic of the Hucumbene and upper Murrumbidgee
Rivers extends into the high plateau about Kiandra, but the treeless Monaro
plains and slopes, the ridge areas and the forested, dissected Umaralla plain give
sharply defined scenic contrasts with abrupt passages from one to another. Five
types may be recognized, as follows:

I

230 THE SURFACE HISTORY OF MONARO, N.S.W.,

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Text-fig. 1.—Locality Map. Part of the Victorian boundary is shown in the

extreme south-west: a heavy broken line represents the Main Divide, lighter

broken lines show other significant divides, and the Federal Capital Territory
is shaded.

1. The High Plateau.—The sources of the Murrumbidgee and Hucumbene
Rivers are comprised in this division: the elevation is consistently great, from
4,000 to 5,800 feet above sea-level, the hills are dome or hummock shaped, except
where weathering basalt flows give rectangular outlines, and they are separated
by wide valleys of gently concave section. The streams are widely spaced, with
extensive marshes and swamps at all levels, up to 2 miles in width, and the
Murrumbidgee above Rule’s Point glides over black, peaty soil that has accumulated

BY IF. A. CRAFT. 2oL

as the result of a prolific growth of shrubs and swamp grasses. The main
physical control appears to be climate: the rainfall is heavy (63 inches at Kiandra),
frosts occur during 9 or 10 months of each year, and exert a powerful disruptive
influence over the whole landscape. Rainstorms sweep away the weathered rock,
and both minor surface features and streams tend to be eliminated.

2. The Basin Country.—Passing eastward of Kiandra, there is a general fall
except in the peaks to the north of the Murrumbidgee, and with this fall comes
the subordination of certain features of the high plateau and the disappearance
of others, such as the swampy valley plains with their accumulations of peaty
clay. Rounded forms are the expression of normally weathering granite and fissile
slate rather than the dominating features, and the breaks of slope between
upland and hillside, and between hillside and valley floor are sharp and distinct
(see Plate ix). The courses of the main streams show alternate basins and
narrows: the Murrumbidgee flows from its headwater plain across a hard strike-
ridge to Kelly’s Plain, whence it passes to Yaouk by a narrow gorge (“the Gulf’),
thence through a rather dissected level to Bolairo plain, and the narrower valleys
and gorges leading to Bredbo. With the Eucumbene below Kiandra, the wider
sections of the gorge do not assume a plain form until Alpine Hill is passed,
where there is a sudden expansion and flats which are partly built up of coarse
river drift. A further narrow place is met before the undulating Adaminaby plain
is reached, beyond which are the Jindabyne levels, a gorge as Beloka ridge is
crossed (quartzites, etc.), and the great terraced valley at Dalgety.

It will be seen that the “basin and narrow” topography is normal in this
section, and there are abrupt passages from level plains and mature forms to
steep ridges and canyons, with some steepening of the stream gradients: there
are considerable alluvial deposits of both coarse and fine material in the basins,
with an apparent maximum thickness of 100 feet at Yaouk, and the streams swing
about in the plains. A general balance seems to have been established between
erosive forces and the resistance of various parts of the country, with rather
steeper river gradients on the harder rocks, and uniform cutting in most places,
the exceptions being the head plain of the Murrumbidgee and the gorges above
Alpine Hill, with lesser and greater downcutting respectively.

3. The Treeless (Monaro) Plain.—A considerable part of Monaro and the
valley of the Snowy has the appearance of a plain when viewed from a distance:
there are extensive areas of level country about the Main Divide, but elsewhere
the original surface has been lowered to give a series of inclined plains which
have been trenched with narrow valleys. In places the development of horizontal
levels is favoured by the presence of basalt flows which filled valleys to the level
of the ridge crests, and even built up the divide south of Cooma, eventually
giving a treeless plain, now apparently extended westward by the clearing of
granite country of a similar elevation. The valleys of the granite section are
wide, terraced, and gently undulating with convex slopes, but the basalts have a
considerable number of lagoons on their higher parts, with accompanying indefinite
drainage, and sharp breaks of slope where various flows have been exposed by
weathering and erosion.

The general rise to Kosciusko and the high plateau on the west of the
Eucumbene-Snowy line is marked by the presence of narrow, steep sided valleys,
whose contrast with the gentler and lower Monaro levels may be partly due
to rapid cutting of deep valleys along cardinal lines of weakness, the quick
removal of weathered material, and the absence of an effective layer of decomposed

232 THE SURFACE HISTORY OF MONARO, N.S.W.,

rock to hold water and promote weathering. On the east, conditions appear
to have allowed slower stream cutting, with concomitant gentler slopes, more
effective weathering, and eventual general reduction. Thus the cutting streams
of the Berridale and Dalgety districts are exposing well rotted granites in their
beds, in areas where the lowering near watercourses has not been disproportionate
to that of the channels themselves, and they contrast with the more powerful
streams of the gorge country, whieh run over fresh rock as a general rule. In
the granite peaks of the Federal Territory and Yaouk, rapid valley cutting may
have assisted in the preservation of the high points, but outward-sloping separation
planes also help, and small differences in chemical composition may be found
to have a considerable effect on the rate of weathering.

4. The Forested (Umaralla) Plain.—The forested slopes of the Umaralla
drainage contrast with the treeless plain, although the two are of a similar
elevation. With the former, the high-level plain has been dissected by streams
that fall from the swamps and undulations of its surface into steep-sided
valleys, in which they have gentle grades through most of their winding courses.
When the valleys approach the Murrumbidgee they are broad and flat-bottomed,
with falling dividing ridges on either side of each, and a generally asymmetric
cross-section: proceeding upstream, the ridge outlines become more angular as
the valley sides close in, and dissection is not apparent on the Tuross divide,
although the eastern extension of the plain into the coastal territory is trenched
with deep gorges. There is higher land both towards Gourock and Nimmitabel,
led up to by forested ridges.

5. The Ridge Country (northerly Murrumbidgee)—The features of this
unique area are being described by Browne, who emphasizes the dependence of
its topography on the existence of parallel zones of rock with great differences
of resistance to weathering and erosion. Standing rather away from the river
is the ridge country of Yaouk Hill and the basins of Bredbo and Queanbeyan
Rivers, where the higher points fall gradually in a series of branching ridges,
and where the slopes of dissected terraces take on a like appearance owing
to the establishment of closely-spaced tributary streams which have not been
subjected to a process of integration.

Physiography.

While the appearance of the country varies regularly from place to place,
there is also a definite order in the suecession of erosional features and levels
that have been moulded to give existing forms. Sussmilch (1909) recognized a
surface of reference which he termed the “Monaro peneplain’’, that was differenti-
ated by warping and faulting to give horizontal or tilted blocks at various
altitudes. Thus the Monaro peneplain at 3,300 feet in the Cooma district was
conceived to have been co-extensive with the Yass-Canberra plain to the north,
now at 2,000 feet, with a tilted surface developed between them along the
Murrumbidgee. However, the hill crests and the benches rising to 3,000 feet or
rather more along the course of the river could also be interpreted as lying in
an extension of the high-level peneplain about Cooma, with the Yass-Canberra plain
appearing as a much newer feature. Browne’s work shows the probability of this
explanation being correct, and a comparison might also be made with the neigh-
bouring Shoalhaven Valley; the Monaro peneplain extends across the Umaralla
basin and has a considerable development in the Shoalhaven area (Craft, 1932a),
with a further extension into the Molonglo drainage to the north-west, and into
the Canberra district, a lower peneplain at 2,000 feet (the “Shoalhaven Plain’’)

BY F. A. CRAFT. 233

being directly comparable with the Yass-Canberra plain. The extent of erosion
in Monaro since the development of the main levels is shown by Tertiary basalt
flows, which may be considered briefly under two heads: those of Monaro and
Kiandra.

The Monaro Basalts.

The undulating surface of Monaro peneplain falls from 3,200 feet near the
Cooma-Berridale road to 3,000 feet towards Cooma, and to a slightly lower
elevation near the Snowy at Dalgety. Narrow valleys were eroded in it to a
depth of 500 feet as an extension of the lower peneplain (= Yass-Canberra or
Shoalhaven plain), and were filled) with basalt that buried stream gravels at
various altitudes, the highest being at 3,750 feet on Dry Plain; basalt in that
vicinity rises just over 4,000 feet, and to a similar elevation on the main divide
near Nimmitabel. Much of the volcanic covering has been removed, so that
considerable parts of the pre-basaltic surfaces are exposed in the Cooma district,
and notable terraces have been formed in the high survivals between Cooma and
Bombala. Where stream erosion has been most active in attacking the built-up
surface, the buried valleys have been partly exposed with remnants of basalt
adhering to their sides. Browne has noticed this feature to the north-west
of Cooma, and it is also seen in the valley of Bobundara Creek near Maffra,
while the valley of Goorudee Rivulet near Bolairo shows contact quartzite from
the basalt contacts in situ between 3,250 and 3,500 feet; this latter overlies pebbles
and white clay, and coincides with the modern valley slopes down to stream
level (Plate ix).

Three stages are thus revealed: the general weathering and stripping of
basalt sheets from the Monaro peneplain, the cutting of valleys through the
basalt and their extension by headward erosion, and a process by which the
basalt was removed from the filled valleys until, with the re-establishment of
the former slopes, erosion has virtually ceased. The only outstanding example
of the last-named is in a basin where old-age features were not being subjected
to considerable erosive attack, and doubtless chemical weathering of the basalts
was the main factor involved in their final destruction.

Kiandra Basalt and Lead (Text-fig. 2).—The greater part of the highland
basalt of Kiandra covers a deep lead of stream drift and peaty clay (‘‘lignite’’
of miners), with an extreme vertical range of 1,600 feet from the basalt at
Rule’s Point (4,250 feet) to Tabletop. Andrews (1901) showed that the main lead
is 150 to 170 feet thick, and that it occurs in a rock-bound channel apparently
falling northward across the present Eucumbene divide to the Murrumbidgee: on
the southern bank of that river the drift is 50 feet thick, but this decreases to a
few inches on the opposite side. The leads were covered with flows of basalt
up to 300 feet in thickness, which form a capping for the monadnocks of Tabletop
(5,850 feet) and Selwyn (5,288 feet), with a general northward fall of both
upper surface and base, and an upward limit of 5,300 feet over the leads themselves.
The present valley of the Eucumbene at Kiandra, with a depth of 600 feet, is
clearly a post-basaltic feature, and gives a measure of active erosion in this
section of high rainfall that contrasts with the static conditions noticed at
Bolairo.

Andrews’ sections show that the lead is not dissimilar to other upper
Tertiary leads in this State, as the drift lies below the general level of the
neighbouring plateau on either side, and the basalt has ceme up to this surface,
although the specially active erosion of the district has removed parts of the

234 THE SURFACE HISTORY OF MONARO, N.S.W.,

original containing walls on either side. Sussmilch and Jensen (1909) and
Andrews (1910) classified the flows with the former’s “monadnock basalts”, but
in view of the position of the greater part below the general level of the plateau,
this can hardly be sustained without definite fossil evidence. The distinctive

| Murrumbidgee Rj

Gooandra

fo00Ft, ViH= 4).

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aa
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S|
al
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Text-fig. 2.—Stream change in Kiandra district (after E. C. Andrews), also
profile of lead channel, with basalt in plain black, and drift barred (drawn
from details from E. C. Andrews).

feature of the lead is the presence of peaty clay, but this only differs from
the light clays of such leads as the Shoalhaven by as much as the peaty deposits
of the modern upper Murrumbidgee differ from the light-coloured flood-terraces
of the Shoalhaven.

The origin of the leads is indicated by Andrews’ levels, which show the profile
of their channel base to be a normal curve of erosion, with no apparent dis-
continuities (the section of low grade at Kiandra is duplicated in part of the
Eucumbene channel), and the lead deposits have a notably uniform thickness
on both steeper and gentler grades. Allowing for normal variation, they show
considerable uniformity in the basal ‘gutter wash’, the lower horizon of peaty
clay, and the sandy beds which predominate towards the upper levels. There
appears to have been simultaneous deposition along the whole channel length,
which may have been due to changes in the load and volume of the transporting
stream, or to some violent interruption in its history: it has been suggested
that such deposits were due to widespread subsidence bringing the country below
sea-level (Wilkinson, 1882; Andrews, 1910), or to a blockage by basalt flows
in one case (Craft, 1931), while warping and faulting, decreased rainfall or
increasing maturity have also been mentioned as possibilities (Andrews, 1915), but
none of these applies to Kiandra. The normal profile of the channel base has
been noticed: it has not suffered notable warping or tilting, as the general level
of the plateau at 5,000 feet or a little more extends northward from the head

te
i)
or

BY F. A. CRAFT.

of the lead to the headwaters of the Murrumbidgee, and there is no evidence
of warping or faulting across the channel, or of blockage by basalt flows, which
came later to cover the drifts. Likewise, there is no suggestion that the northerly
country rose across the stream course to give conditions of still-water deposition
at any stage, because the channel at Rule’s Point is based on a peneplain level
which included the plain divide at the head of the Murrumbidgee, only 130 feet
above the lowest point of the drifted channel. The basalt extends on to this
plain at Rule’s Point, virtually clear of the drifted area: deposition as the result
of subsidence cannot be admitted with deposits restricted to a definite channel,
even though preservation on plains was made possible by basalt flows, and where
they are of a reasonably uniform thickness at all grades, showing no great angular
unconformity with their channel.
There is no alternative but to conclude that the drifts were laid down as
the result of changes in the transporting stream itself which substituted deposition
for erosion. Moreover, since the change was uniform along the whole channel,
variation as the result of such external action as stream capture is unlikely, and
the then-existing stream simply became incapable of gaining large fragments from
the landscape, and of transporting all the material brought to the main channel:
this may indicate an increased cover of vegetation and a decrease in the effective
power of the flood stream. Modern conditions in the upper Murrumbidgee show
that peaty deposits accumulate at grades as steep as 1:50, and with streams
of less volume earrying more drift from weathering hillsides, it is probable
that the thickness of peaty material would continue to increase until changing
conditions altered the character of succeeding deposits, or renewed channel
cutting. Another resemblance between modern conditions and those of the past
is also found at the head of the Murrumbidgee, where the peaty layers are
stratified, and contain bands of leached clay in places.

Summarizing, it is found that the drift of Kiandra lead was accumulated
as the result of changes in vegetation and hydrography, without tectonic dis-
turbance or the blocking of the stream channel. Some of the material was deposited
at a grade of 1:65, which compares with grades of approximately 1:50 in
steeper parts of the modern peaty clays, 1: 58:5 of the Vegetable Creek lead
(David, 1887), and 1:60 which may be observed in many existing stream
terraces: the figures give some idea of the limiting slopes in such cases. Many
of the leads of eastern New South Wales probably have a similar origin to
that of Kiandra, although it may still be necessary to explain some by tectonic
or basaltic interference. Furthermore, it will be seen that the basalts of Monaro
and Kiandra give no evidence of differential uplift since they were poured out;
on the contrary, they are associated with surfaces which bore the same relation
to one another as they do now—a condition which gives a great measure of
freedom in the interpretation of old erosional levels, for which Monaro peneplain
may be used as a standard.

Monaro Peneplain.—A consideration of the upper Murrumbidgee shows that
post-basaltic erosion has tended to reproduce pre-basaltic features in detail, with
some downcutting in the Cooma district and in the various gorge and basin
sections between Rule’s Point and Bolairo; in general, the lowering due to
this action is between 100 and 200 feet, and the lack of great local stream
revivals shows that post-basaltic warping or faulting has been a minor factor in
the development of surface features, a conclusion that applies equally to the
Eucumbene-Snowy line between Kiandra and Maffra, with downcutting in excess

236 THE SURFACE HISTORY OF MONARO, N.S.W.,

of 200 feet in the gorges above Alpine Hill, and greater valley formation at
Kiandra. The main levels and terraces are thus older than the basalts, but
their relationship to one another is not immediately evident, since correlation on
the basis of height only is risky, and the meeting of surfaces is not infrequently
obscured by other erosional features. The eastern portion is the simplest in
this regard.

Eastern Monaro.—The Shoalhaven Valley shows that a pre-Tertiary peneplain
exists at 3,000 to 3,400 feet above sea-level, and continues into the Umaralla basin:
there is a normal rise to 4,000 feet between the two systems, and the peneplain
may have been carved out of a master level about this relative position. This
gives the status of the great peneplain of eastern Monaro, with high points
rising to 4,300 feet in the level-bedded Devonian rocks to the east, and an
extension from the Tuross divide to Berridale at 3,000 to 3,400 feet. To the south-
west of Cooma and in the adjoining parts of the Snowy basin, this peneplain
gives a perfectly even skyline that hardly shows the gentle undulations of its
high points: along the northerly Murrumbidgee it is represented by extensive
benches and hills with crests lying in a uniform summit plane between higher
ridge masses, and by passes in the latter features, but towards Canberra its
nature is essentially residual on account of its nearness to the effective edge of
the highlands. In the opposite direction, to the south of Monaro ridge, the high
points rise to a normal height of 3,200 to 3,500 feet, with the highest members
on hard ridges or somewhat removed from converging streams, and with
isolated residual points rising to a maximum of 4,700 feet on the Victorian
border. There is a general lowering to 2,900 feet where the Maclaughlin River
approaches the Snowy, and the main streams flow in broad terraced valleys some
400 to 500 feet lower, although they are only just cutting below the pre-basaltic
levels. It is clear that there has been no effective Tertiary peneplanation here.

Thus the normal features of eastern Monaro may be described as a peneplain
above 3,000 feet, with residual points rising 1,000 feet higher in the east, and
the larger masses of Gourock and Tinderry attaining maximum elevations of
4,800 and 5,300 feet respectively, with uniform ridge lines about 4,600 feet, and
a considerable further development to the west of the Murrumbidgee. It has
already been concluded that Gourock Range was thrust above the Shoalhaven
Valley before the formation of the Monaro peneplain (Craft, 1932a), and from
what has been written above respecting surfaces in extra-Shoalhaven areas, it
might be inferred that the high block now above 5,000 feet extended over eastern
Monaro, where it has been planed down by erosion, leaving the modern high points
on the Tuross divide in a superior position, although they were on the downthrow
side of the fault. Western Monaro proves instructive in this respect.

Western Monaro—tThe high plateau between the Kosciusko block and the
head of the Murrumbidgee extends over a meridional distance of 50 miles at an
altitude of 5,000 feet or a little more, and has a general width of 8 to 10 miles
except in the vicinity of Jagungal (monadnock), where it is considerably wider,
although furrowed by deep gorges. The greater part of this surface presents an
even skyline with residuals rising to 500 or 600 feet above it, and to 1,500 feet
in the single case of Jagungal, but the extension into the Murrumbidgee drainage
has been maturely dissected, although the hills and ridges rise to a uniform
summit plane. So far as the more northerly part is concerned, the terraces
cut by the Eucumbene west of Alpine Hill and occurring between 4,400 and 4,600
feet are quite extensive, and penetrate to a depth of a mile or two from the

bo

~]

BY F. A. CRAFT.

river in areas of softer rocks, although they are less perfect where the stream
cuts across hard meridional bars, and they have been much dissected by gorge
cutting. On the other hand, a very perfect set of plain levels is developed between
4,200 and 4,400 feet at the heads of the Murrumbidgee and Goodradighbee, forming
a plain which comprises stream valley and divide indifferently, and encircles such
high plateau ridges as Nattung (5,300 feet), where there are occasional passes
about 4,500 feet. The plain is subordinated in the high ridges crossed by the
Murrumbidgee below Rule’s Point, but it is represented by swampy expanses on
the courses of Nungar and Tantangara Creeks, separating Nungar (5,608) and
Gang Gang (5,321) hills from the high country about Kiandra. The two surfaces
are well defined with respect to one another: the lower plain at 4,200 to 4,400
feet has resulted from the dissection of the higher level above 5,000 feet, whose
residuals become more scattered in the direction of Adaminaby, but whose ridges
are prominent in the southern half of the Federal Territory.

Perhaps the most perfect development of the lower of these surfaces is found
to the east of Kelly’s Plain, where the Murrumbidgee crosses it in a gorge
(Plate ix), and although the apparent eastward limit of the section is at
Yaouk, foothills of the higher residual points about Yaouk Plain rise to 4,300
feet, and the gap at the head of the Cotter River is a little higher. The relationship
of the plain above 4,200 feet to such high points as Yaouk, Morgan and Bimberi
is clearly defined: it has been cut into their original mass, making them comparable
with the high plateau immediately to the west, the superior elevation of a
few points being doubtless due to their inherent resistance to erosion.

Passing eastward along Monaro ridge, the higher points are no longer found
beyond Adaminaby, but the master surface is an extension of the lower level
we have just been studying, and the general elevation of the divide is over
4,000 feet to Rhine Falls: hills and ridges come to that height on the Murrumbidgee
side to give a fairly uniform skyline, while the area in the Rhine Falls-
Jindabyne-Adaminaby triangle is slightly higher, with an exceptional point rising
to 4,816 feet. The level is again found on the flanks of Yaouk Hill and to the
east of Alum Creek, with a fall of varying steepness to the river.

Considering the eastward limit of this high plain, we find an undulating
slope to the Monaro peneplain at 3,000 feet within the salient of the Murrumbidgee
near Cooma; there is a sharper fall to the west of Peak Creek and about Rhine
Falls, through 800 feet in places, but the impressiveness of the feature is lost
when the Main Divide is passed, and the south-easterly slope in the drainage area
of Wullwye Creek is in a series of irregular undulations over a distance of some
miles to the Berridale district. But there are extensions of the Monaro levels
behind this eastern face, both along the two main rivers and creeks which have
cut back from them, the Murrumbidgee examples being very impressive.

The topography of the basins of Kelly’s Plain (3,800 feet), Yaouk (3,600 and
3,700 feet) and Bolairo (3,200 feet) is essentially pre-basaltic, since there is clear
evidence of basaltic visitation in the valleys of Bolairo, 600 feet below Monaro
ridge, and the other basins also lie below that horizon. A similar inference
could be drawn from the relative positions of the Rule’s Point basalt (down to
4,250 feet) and the main summit plane between Kelly’s Plain and Yaouk, which
is about 4,400 feet, and also from the form of Yaouk Plain itself, which is cut
across the grain of the ceuntry with a perfection not associated with any definitely
post-basaltic features anywhere in the region. Yet these basin plains are young
when compared with the surface in which they are eroded, since they are

238 THE SURFACE HISTORY OF MONARO, N.S.W.,

separated by a gorge section through hard quartzites, etc., above Yaouk, while
the superior level about 4,400 feet was formed by the bevelling and planation of
those hard folded strata, even across the divides—a process that must have
occupied a great time, and indicates a corresponding age for the surface concerned.

The position becomes more complex to the east of Adaminaby and Bolairo,
for gently shelving plains make their appearance and fall towards the Murrum-
bidgee along the established creeks: the initial slope from Monaro ridge is steep
for 400 or 500 feet down to 3,600 or 3,700 feet above sea-level (up to 30 degrees),
whence a gentle grade is assumed through a vertical distance of 200 or 300 feet
to a steeper slope towards the river. Ridges and residuals rise close on 4,000 feet
(Plate ix), and there is a repetition of the features in the great salient of the
river on the opposite bank: the undulating plains of Bolairo are associated with
powerful streams and a zone of low resistance, but relics of the higher landscape
persist eastward along the right bank of the Murrumbidgee until the converging
streams of the Cooma district are reached. The features of the Murrumbidgee
are repeated in the Eucumbene drainage, with somewhat different forms owing
to a greater proportion of granite on the landscape: this is especially the case
in the Berridale-Dalgety section, but along the eastern side of the EHucumbene
between Adaminaby and Jindabyne the high level plains at 3,800 to 3,900 feet
break sharply to the uniform valley wall, with a fall of the order of 500 feet
to the river, rounded forms being characteristic of higher points and of minor
features. There are limited plains along the course of the main stream, the
principal being in front of Alpine Hill (3,700 feet), Adaminaby (3,500 feet) and
at Jindabyne (2,800 feet), the second being the most notable because it extends
up the broad valley of Fryingpan Creek, a stream flowing against the general
run of others.

From these facts, it seems that the plain which rises from 4,100 feet at Rhine
Falls to 4,400 feet west of Adaminaby is the main feature of this part of Monaro;
that it was cut in a higher mass that still occurs in a much reduced form
to the north and west, including the highest land on either side of the northerly
Murrumbidgee. Further erosion of the features above 4,000 feet has caused a
general lowering towards the east, and a selective cutting of local plains along
the courses of the main streams down to the level of the Monaro peneplain,
which may be coeval with them, despite its great extent and level character
over eastern Monaro. The surface variety which Browne describes along the
northerly Murrumbidgee lends support to this contention, because it shows how
sudden a change is possible from low to high relief with the geological variation
found in the region. It would appear that the only place where block-faulting has
been indicated as a comparatively recent possibility is in the Kosciusko plateau
which rises above 6,000 feet, and the probable fault indicated by David (1908) as
throwing 200 feet at Pretty Point is some indication of the magnitude of individual
displacements to be expected towards the limits of that restricted area. It is not
suggested that warping and faulting did not occur elsewhere on the tablelands
which have been described, but that such movements were of a minor character
and had comparatively little influence on the surface features, which are essen-
tially due to the erosion of a mass that has been subjected to fairly uniform vertical
movements since closing Palaeozoic time. This idea accords with the stream
relationships.

Streams of Monaro.—The first account of the streams of the region is found
in the narrative of W. B. Clarke, who correlated the streams having meridional

BY F. A. CRAFT. 239

tendencies with the grain of the country, folding, and the occurrence of soft zones,
while the others were recognized as following definite fractures which fall into a
series of families, the principal trending NW-SH, NE-SW, and E-W: rivers of
the Bombala district were ascribed to a surface depression and contrasted with
those radiating from Kosciusko, a point of maximum uplift. David (1908) added
late Tertiary block-faulting about Kosciusko as a causative factor in the develop-
ment of the present shape of the Crackenback and Snowy, while Sussmilch (1909)
developed a similar idea with respect to the northerly Murrumbidgee, which he
conceived to have come into existence as the result of warping and faulting of
a peneplain surface raised unevenly to form the present highlands. He believed
that the upper Murrumbidgee and the southern part of its meridional course
originally discharged to the Snowy, thus following Clarke in principle, who had
described Monaro ridge as an anticlinal feature, and had referred to great stream
dislocations as the result of basalt flows. Griffith Taylor’s opinions with respect
to the Murrumbidgee were rather similar, but he argued more from the evidence
of stream shapes, and held that the tributaries flowing contrary to the general
northerly direction of the Murrumbidgee belonged to an earlier southward flowing
system. Doubt was cast on some aspects of these views by Browne (1928), who
described the valley of the northerly Murrumbidgee as a non-tectonic feature,
and by Craft’s (1932a) demonstration of the parallel Shoalhaven area as one that
had suffered no appreciable differential uplift from north to south over a period
antedating the basalt flows and coming to the present time, and whose features
extend normally into the Murrumbidgee basin.

The correlation of the stream lines with a definite fracture pattern is sound,
and requires no further elaboration, but they may also be classified by reference
to the stage of development of their various networks, and may be grouped under
three headings, namely: (i) Those which have failed to develop a complex
system of tributaries: the Cotter, Gudgenby, Naas, much of the upper Snowy
and of the northerly Murrumbidgee are included. (ii) Those which present a
minutely branching system, such as Queanbeyan and Bredbo Rivers and Alum
Creek. (iii) Those which have integrated their tributaries into a few large,
evenly spaced branches: this class is of particular importance, since it comprises
the Murrumbidgee and Hucumbene-Snowy systems on either side of Monaro ridge.

Of these three, the first class has been most powerfully influenced by geological
structure: the Cotter, Naas and northerly Murrumbidgee flow along zones of weak-
ness flanked by persistently hard ridges, while the Gudgenby and upper Snowy are
marked by linear courses, clean angular junctions, and restriction to well defined
fissures, suggesting development along these to the general detriment of the
pattern, and a lack of closely spaced minor weaknesses to give rise to smaller
streams. In the second class, each small tributary flows in a distinctive valley
of its own, and there is a correlation with high ridgy or terraced country which
the multitude of streams is unable to reduce efficiently.

The third class is of a composite nature, and its various members have
developed under a variety of circumstances. West of Yaouk and Adaminaby the
simplification of patterns appears to have been effected by heavy rainfall on a
weathering landscape, and the identity of some of the existing streams is almost
lost where they flow across level swamps. Hast of Adaminaby, the minor valleys
have been greatly widened and are frequently of a trough shape, with a degenera-
tion to broad, shallow hollows with only a few definite watercourses to collect the

240 THE SURFACE HISTORY OF MONARO, N.S.W.,

run-off from the gentle surfaces: this is the case in both granite and basalt areas,
whose streams are remarkably similar to one another in plan, despite the fact
that hundreds of feet of basalt have been removed from parts of the Monaro
peneplain while other rocks have suffered comparatively little denudation. In
the Umaralla drainage, the present stream system appears to have been inherited
in detail from the Monaro peneplain at 3,300 feet, although not much of it now
flows over that surface: dissection has not caused a complex system of tributaries
to develop on the hillsides and ridges as in the country immediately to the north,
possibly on account of insufficient rainfall (the rainfall at Cooma, under rather
similar conditions, averages 19 inches per annum). Thus the great feature of
Monaro is the presence of a genetic stream type both on the more ancient surfaces,
and in places affected by basalt flows; the spacing and general arrangement of
tributaries shows considerable uniformity, and it is not a fortuitous circumstance
that the distribution of the three types should be ordered as it is: the first two are
associated with lands that persist at a relatively high elevation, while the third
is virtually confined to those parts which have been susceptible to peneplanation
in the more distant past, and to elimination of surplus tributaries in more recent
times.

Stream Origins. (Text-figs. 3 and 4.)

The Monaro region is notabie as the place where the Main Divide of New
South Wales makes a westward turn into Victoria. Perhaps it is this circum-
stance allied with the general flatness of the continent which has led to the
highlands about Kosciusko being described as an Alpine knot, but the term is
hardly applicable, as the plateau is a relatively simple surface crossed by low
erosional ridges, which are not obscured by possible block faulting about Kosciusko
itself and in the neighbouring highlands of Victoria, although deep gorges give
a complex topography in places. There is no reason why the co-extensive plateaus
and drainage systems in the two States should be considered apart from one
another, and they have one salient drainage feature in common in the presence
of meridional streams discharging northward to the Murray or Murrumbidgee,
or southward to Bass Strait (Text-fig. 3). Apart from the fringe of Pacific coastal
streams, the Main Divide is a simple affair with opposed salients at the heads
of the Snowy and Mitta Rivers, and a dominant east-west trend. It has been
indicated that the Shoalhaven-Murrumbidgee family may be descendants of ancient
longitudinal streams flowing northward to the Permo-Triassic depression, a sug-
gestion which is in line with the evidence cited by David (1911), and others
whom he quotes, to indicate that the Victorian Main Divide was essentially
developed at the close of the Palaeozoic or the beginning of the Mesozoic era. The
streams of the region thus originated parallel to the ancient grain of the country,
and have had their expansion confined by zones of hard rock or of specially
resistant features: it is only in Monaro, to the south of the geologically varied
Federal Territory-Gourock mass that a wide area of more uniform rocks has
allowed the ready development of peneplain features, and the formation of
more extensive stream systems. The straight eastern divide shows the uniformity
of attack from the Pacific or, perhaps, the uniform resistance offered by meridional
trends, and the modern piercing of the divide to the east of the Shoalhaven is
readily explained in terms of nearness to the coast and the strong attack by
revived streams along definite lines of weakness.

BY F. A. CRAFT. 241

Tin Aq

15O°= ——— :
Gee Oe eo MILES
Text-fig. 3.—General Stream Pattern of the extreme south-east of Australia.

The Main Divide is shown by a broken line, and the principal streams are:
1, Murrumbidgee; 2, Murray; 3, Mitta; 4, Mitchell; 5, Snowy; 6, Shoalhaven.

As Clarke suggested, it is possible that the eastern Snowy drainage was
originally directed towards a depression discharging westward to the lower
Snowy, but the present lowering of surfaces is to be attributed to erosion because
the high points lie in a definite summit plane. In plan; the pattern suggests
development from a main east-west tributary of the Snowy through Bombala,
with an overgrown branch extending towards the north-west. It might he sug-

242 THE SURFACE HISTORY OF MONARO, N.S.W.,

gested, aS an interesting speculation, that the upper Murrumbidgee, which is
placed roughly between the Federal Territory granites and those of Dalgety-
Kosciusko, can only be explained in some such manner, and although it cuts
across granite lines in places and is well adjusted to structure, it may have
originated in an old high-level depression that persisted from the time of the last
folding movements and granite intrusions to which the region was subjected: the
possibility of such an explanation being feasible is shown by the stability of the
streams as far back as definite physiographic records go.

Sussmilch (1909) and Taylor (1910) have suggested that the upper Murrum-
bidgee originally flowed to the Snowy, whence it was diverted northward by
capture following warping and basalt flows. The balance of both pre- and post-
basaltic erosional features on either side of the present Main Divide does not
favour the idea, and there is no certain evidence of considerable warping. More-
over, the Umaralla appears to have discharged northward over a very long period

pects Mbidzee
Sagres Snowy

Text-fig. 4.—Profiles (Talwege) of the Murrumbidgee and Eucumbene-Snowy
Rivers. Vertical exaggeration = 50.

of time, and there are no features rising above the level of the Monaro peneplain
between it and the Murrumbidgee, and no suggestion of an old divide. Considering
the profiles of the main streams (Text-fig. 4), it will be seen that they are
very similar over the upper parts of the courses, in which the streams flow on
opposite sides of Monaro ridge, although the revived lower part of the Snowy
has a steeper fall to the nearer coast. Simultaneous development to a comparable
stage is shown for the two rivers, and the probability of a major capture in
their history of the nature suggested is exceedingly remote. It is true that the
Eucumbene has established new courses about Kiandra, but this seems to give
a measure of the change of the two systems with respect to one another.
The existing divide has been an essentially stable feature over a long period
of time, antedating the basalt flows considerably.

SUMMARY.

The Monaro region appears to have been elevated above the country to the
east, probably in closing Palaeozoic or early Mesozoic times, after the reduction
of the dominant Kanimbla (Hercynian) features. Since then, it appears to
have been subjected to a series of epeirogenic uplifts which induced widespread
peneplanation, of decreasing completeness to the west and north. Relics of the
original surface at 5,000 feet or more compare with the level-bedded Devoniai
rocks towards the eastern divide about 4,000 feet, and peneplain levels exist
at 4,000 to 4,400 feet and about 3,200 feet respectively, the latter (Monaro
peneplain) being comparable with the older features of the Shoalhaven Valley.
The development of steeper-sided valleys towards the east was proceeding before
the outpourings of the (‘‘newer”’ or upper Tertiary) basalts, and more recent

Proc. Linn. Soc, N.S.W., 1933. PLATE IX.

eae

Monaro District, N.S.W.

me

a

t

BY F. A. CRAFT. 243

erosion has been largely devoted to the removal of basalt and the re-development
of the buried valleys. In this connection the high Kiandra lead is held to be
a product of stream action without tectonic interference, and its district contrasts
with the lower country of eastern Monaro in which the buried stream drift is
of relatively small extent and thickness. Typical land forms have developed in
response to climatic and geological factors, giving rounded features in the
high plateau of the west, a “basin and narrow” topography] on the edge of
the higher blocks, and a dissected plain over much of the rest of the area. The
stream history is one of stability over long periods of time, and a correlation
is found between the modern stage of pattern development and the past sus-
ceptibility of the various sections to peneplanation. It is held that the stream
system is inseparable from that of eastern Victoria, and that the significant divide
is a more or less easterly continuation of the Main Divide of Victoria, with the
meridional portion north of Nimmitabel having little tectonic significance. It
might also be noted that the eastward fall of erosional surfaces begins definitely
some 8 miles east of the Umaralla-Snowy divide in the area discussed, and
although the probability of post-basaltic warping and faulting during the most
recent uplifts is not excluded, it is believed to have had little influence on the
development of the modern surface.

References.
ANDREWS, E. C., 1901.—Report on the Kiandra Lead. Geol. Surv. N.S.W., Mineral
Resources, No. 10.
, 1910.—Geographical Unity of Eastern Australia. Journ. Roy. Soc. N.S.W.,
xliv, 420.
, 1915.—Geological Survey of the Cargo Gold Field. Geol. Surv. N.S.W., Mineral
Resources, No. 19.
BROWNE, W. R., 1928.—On some Aspects of Differential Hrosion. Journ. Roy. Soc.
INES Wise Xe ide
(Paper on Cooma district and the northerly Murrumbidgee in preparation).
CLARKE, W. B., 1860.—Researches in the Southern Gold Fields of N.S.W., Sydney.
CraFt, F. A., 1931.—Physiography of the Shoalhaven River Valley. iv. Nerriga. Proc.
LINN. Soc. N.S.W., Ivi, 412.
, 1932a.—Idem. vi. Conclusion. Proc. LINN. Soc. N.S.W., lvii, 245.
, 1932b.—Notes on Erosional Processes and Stream Gravels. Proc. LINN. Soo.
N.S.W., lvii, 280.
Davip, T. W. E., 1887.—Geology of the Vegetable Creek Tin-Mining Field. Mem. Geol.
Surv. N.S.W., Geology, No. 1.
, 1908.—Geological Notes on Kosciusko. Pt. ii. Proc. LINN. Soc. N.S.W., xxxili,

Gibie
, 1911.—Notes on some of the Chief Tectonic Lines of Australia. Journ. Roy.

Soc. N.S.W., xlv, 4.

SUSSMILCH, C. A., 1909.—Notes on the Physiography of the Southern Tableland of N.S.W.
Journ. Roy. Soc. N.S.W., xliii, 331.

SUSSMILCH, C. A., and JENSEN, H. I., 1909.—Geology of the Canobolas Mountains. Proc.
LINN. Soc. N.S.W., xxxiv, 157.

Taytor, T. G., 1910.—Physiography of the Proposed Federal Territory at Canberra.
Comm. Bureau of Meteorology, Bulletin No. 6.

WILKINSON, C. S., 1882.—Notes on the Geology of N.S.W., Dept. of Mines, Sydney.

For an account of part of the Geology, see Browngr, W. R., 1914.—Geology of the
Cooma District, N.S.W. Journ. Roy. Soc. N.S.W., xlvii, 194.

For approximate topographic detail, see International Map of the World (1:1,000,000),
Sheets SI55 (Canberra) and SJ55 (Melbourne).

EXPLANATION OF PLATE IX.
Topography of Western Monaro. ;
1.—Head of Murrumbidgee River near Peppercorn Hill (left); the valley divide of
Goodradigbee River is in the middle distance, behind the black line of peaty swamps
along the river.

244 THE SURFACE HISTORY OF MONARO, N.S.W.

2.—Murrumbidgee River at Kelly’s Plain, looking eastward. The river cuts across
the high terrace in a gorge to Yaouk (right background),*with the high hills of the
Federal Territory on the extreme left.

3.—Topography of Yaouk Basin, looking eastward, with Yaouk Hill on the right;
this is a “close-up” of the right-hand portion of No. 2.

4.—Murrumbidgee River at Bolairo, looking westward.

5.—View northward towards Bolairo from the Adaminaby-Dry Plain road, to show
surface deposits of probable late Tertiary age (white) and their correspondence with
the modern landscape. The Murrumbidgee flows across the background from left to
right.

6.—Eucumbene River from the Adaminaby-Kiandra road, looking upstream with
Alpine Hill on the right: the level plain consists of coarse gravels overlain by fine drift
and silt.

VEGETATIVE REPRODUCTION IN DROSHRA PELTATA AND
D. AURICULATA.

By Joyce W. Vickery, M.Sc.
(Plate viii; thirty-four Text-figures.)

[Read 26th July, 1933.

Introduction.

Drosera peltata Sm. and Drosera auriculata Backh. are winter herbs common
in the Sydney district. They perennate by means of small, more or less spherical
tubers, situated at the base of underground stems.

These two species belong to the subgenus Hrgaleium Planch. of the
Droseraceae. Bentham (1864) follows the older classification of the Droseraceae,
in which the genus Drosera is divided into two subgenera, Rorella and Ergaleium,
distinguished by means of their types of perennating organs. In the former,
the short stock or stem forms, at its upper end, the winter bud which will
develop in the following year. In Hrgaleiuwm, on the other hand, the plant is
characterized by the production of a tuber at some distance below the surface
of the ground. It is from this tuber that the bud develops which will form the
plant in the following season.

Planchon (1848), in his monograph on the Droseraceae, divides the genus
Drosera into thirteen sections, on account of some anomalous species. Drude
(1891, in Engler and Prantl) divides it into five subgenera. Both authors retain
the section Hrgaleium on account of the characteristic method of vegetative
reproduction shown by all its members.

Relatively little work appears to have been done on this section of the
Droseraceae, and as both D. peltata and D. auriculata are abundant in the Sydney
district, it was thought that a study of the developmental history of their under-
ground parts, and their relationship to their environment, might prove of interest.

According to Ewart (1930), D. peltata and D. auriculata are both found
throughout the Eastern States of Australia, viz., Queensland, New South Wales,
Victoria and Tasmania, and also South Australia. D. auriculata is in addition
recorded from New Zealand.

In the Sydney district, D. peltata and D. auriculata are abundant on the
sandy soils, but both species are also frequently found on the heavier clay soils.
D. peltata is common on the margins of peaty swamps, especially when the
surrounding vegetation is not very dense. Soil moisture appears to be the chief
factor governing their ecological distribution. In well drained areas, such as on
the tops of sandstone ridges, they are only found in the autumn, winter, and
early spring, while the soil still retains considerable moisture, and soon disappear
when the warmer weather causes the surface to dry. They are only plentiful
during the cooler months of the year. D. peltata is usually the first to appear,
in moist sheltered places; D. auriculata occurs rather later, and apparently

Jj

246 VEGETATIVE REPRODUCTION IN DROSERA,

requires an even moister habitat for its best development. The plants are capable
of passing through their active phase in a few weeks, and may flower and fruit
and die down to the ground in a short period if the moisture conditions become
unfavourable.

D. peltata and D. auriculata are very alike in their habit and vegetative
characters. The leafy aerial stem is erect, somewhat flexuose if very long,
averaging about 20 to 30 cm. in height. The leaves at ground level may be
reduced to short or occasionally fairly long, linear scales, or else they may form
a rosette of leaves with orbicular or almost reniform laminae and distinct petioles.
The scattered foliage leaves above these are peltate or broadly crescent-shaped
on slender petioles.

In D. auriculata the foliage is always of a pale green colour, while in
D. peltata the leaves vary from a similar colour to a quite deep red.

The stem has an underground region varying from less than 1 cm. to 9 or
10 cm. in length, and is terminated by a small, more or less spherical tuber. In
D. peltata this is always of a bright red colour; in D. auriculata it is typically
yellowish.

In the investigation here reported, the annual development of the tubers has
been considered, together with the relationship existing between habitat and the
depth of the tuber below the surface of the ground.

An accessory method of vegetative reproduction is also described, both
D. peltata and D. auriculata exhibiting the phenomenon of epiphyllous budding.
These leaf-borne buds have been studied with a view to ascertaining the precise
way in which they originate on the lamina, and the physiological factors
influencing their production.

Tuber Formation.

The morphology of the underground parts has been described in D. gracilis
by Planchon (1848). His observations were based on herbarium and preserved
material, which proved inadequate for the determination of several interesting
points, and he directed attention to a number of questions which he left to
be elucidated by future research on living material.

Diels (1906a) figures the underground parts of D. microphylla, D. hetero-
phylla, D. rosulata, D. squamosa, and D. bulbosa, but gives no description of their
developmental history.

Morrison (1905, 1907) examined the underground parts of D. bulbigena,
D. calycina, D. stolonifera, and D. erythrorhiza. These species have not been
examined by the writer. Since, however, several observations which have been
made on D. peltata and D. auriculata differ considerably from the description
given by Morrison, and since he did not give any details or figures of develop-
mental stages, it has been thought advisable to give an account of the develop-
ment of the underground parts from the time of the germination of the seed.

Lubbock (1892) gives an account of germination in D. rotundifolia and
D. binata in which the cotyledons are epigeal. D. peltata and D. auriculata differ
from these in that the cotyledons are retained within the seed coat.

Rendle (1925) states that in those species of Drosera in which germination
has been studied, the primary root is absent, its place being taken by a protoeorm-
like development of the hypocotyl, bearing long attaching hairs. This protocorm
is a temporary structure, and is later replaced by adventitious roots developed from

BY JOYCE W. VICKERY. 247

the stem. This condition differs entirely from that which obtains in D. peltata
and D. auriculata.

Seeds of both species germinated readily in the laboratory on moist filter
paper in Petri dishes. The seedlings commenced to appear after about 14 days.
Observations on the early stages of germination and on the development of the
seedling were made from these plants, but the development of the tuber in the
young plant was studied from plants found growing under natural conditions.

The radicle emerges from the seed first, and grows downwards to form a
short primary root (Text-fig. 1). This never attains any considerable size and only
functions for the first season in the life of the plant. The plumule appears soon
afterwards (Text-fig. 2), and the developing seedling remains attached to the
seed by the two cotyledons, which never entirely emerge from the testa.

Text-figure 3 shows a later stage in the germination of the seed of D. peltata.
The stem has elongated considerably, and the foliage leaves have unfolded,
exposing the glandular hairs. The leaves are always small during the first
season’s growth, and are orbicular in shape. Root hairs develop on the radicle,
which remains relatively short. In D. peltata a rosette of leaves is not formed
in the seedling, the internodes being relatively long.

After the shoot has ascended for a greater or less distance, a bud (D, Text-
fig. 4) in the axil of one of the lower foliage leaves becomes active, grows out and
turns downwards. This bud forms a whitish coloured descending stolon, or
‘dropper’, which is destined to bear the first perennating tuber at its apex. For
convenience, this descending stolon will be referred to throughout this paper as
the ‘dropper’.

The ‘dropper’ is morphologically a stem, as was pointed out by Planchon
(1848), and bears small scale leaves. In both D. peltata and D. auriculata the
colourless scale leaves present on the ‘dropper’ are very much smaller than those
figured for D. gracilis by Planchon. They may be seen more clearly on older and
larger plants (see Text-figs. 9 and 11). The scale leaves are homologous with
true foliage leaves, and are potentially capable of developing into them, as is
shown by an examination of some unusual specimens to be described later.

The ‘dropper’ may penetrate about 1 cm. into the soil during the first season,
and the first tuber is formed at its end. During this process the growing point
becomes inverted, because of the cessation of growth on one side of the ‘dropper’,
while the other side increases in size. The parenchymatous tissue then becomes
enlarged and packed with food material in the form of starch grains. Fat also
occurs in the epidermis and in a few cells associated with the vascular bundles.

The dormant apical bud which is now directed upwards can be detected with
a lens as a cluster of minute scale leaves surrounding the growing point. Other
scale leaves occur on the surface of the tuber and are clearly seen in section
(Text-fig. 6), but are usually inconspicuous maeroscopically.

A longitudinal section through the apex of a ‘dropper’ (Text-fig. 5) shows the
apical meristem surrounded by leaf initials. A vascular bundle (L.) passes from
the vascular tissue (V) of the stem to the base of each scale leaf, but does not
pass into the leaf tissue itself. A slightly unequal growth on opposite sides of
the stem can already be noticed.

A longitudinal section of a young tuber formed at the apex of a ‘dropper’
(Text-fig. 6) shows the apical bud now pointing directly upwards, and the vascular
bundles forming a loop in the ground tissue. Further enlargement of the ground

248

, cotyledons; X, testa. x 32.

VEGETATIVE REPRODUCTION IN DROSERA,

U
Chis €
=

Text-fig. 1—A very young seedling of D. peltata. R, radicle; P, plumule;
cotyledons; X, testa. x 34.
Text-fig. 2.—A young seedling of D. peltata. R, primary root; P, plumule;

Text-fig. 3.—A seedling of D. peltata. R, root; H, root hairs; X, testa;
aerial stem; B, apical bud. x 9-5.

Text-fig. 4.—A seedling of D. peltata with a ‘dropper’. R, root; X, testa;
‘dropper’ arising in the axil of the foliage leaf (A); F, foliage leaves. x 10.

BY JOYCE W. VICKERY. 249
tissue takes place until the diameter of the tuber is many times that of the
‘dropper’.

Rendle (1925, p. 193) states that in the subgenus Hrgaleium, the ‘bulb’
“eonsists of very closely united leaves of which only the tips are free, and form
a peristome-like crown surrounding the base of the hypogeal caulome’’. I have
not seen this condition in any of the specimens examined. The tuber has
always been found to be a solid mass of ground tissue bearing small scale leaves.

The tuber formed during the first season is very small, frequently only about
1 mm. in diameter. At the end of the growing season, the aerial and underground
stems die, and the tuber remains in the dormant state.

The development of the seedling in D. auriculata follows a similar course.
In a young seedling of D. auriculata (Text-fig. 7) the tips of the cotyledons are
again seen to be retained within the seed coat. At a later stage (Text-fig. 8)
the shoot differs from that in D. peltata in that the leaves are arranged in a
rosette, the internodes remaining very short. The radicle (R) forms a short
unbranched root covered with numerous long root hairs. The ‘dropper’ (D)
arises in the axil of one of the lower leaves. The formation of the tuber at the
end of this ‘dropper’ and the subsequent development of the underground organs
is similar to that in D. peltata.

At the commencement of the next growing season the apical bud of the
tuber grows up through the soil, and on reaching the surface, it forms a rosette
of leaves. The arrangement in a rosette of the first few foliage leaves, almost
invariably takes place during the first few years in the life of any individual
plant in both species. In an older plant this is not always the case. There may
then be a small cluster of scale leaves at ground level (Text-fig. 11). At times
these scale leaves may be quite long (Text-figs. 13, 15). The elongated leafy
stem, at the apex of which the flowers are borne, develops above the rosette.

At this stage it will be convenient to define the terms which will be used in
the description of subsequent development. That part of the stem which develops
from the tuber, and which is situated below the surface of the ground, will be
referred to as the underground stem (the rootstock of Morrison, 1905). The
aerial stem continues the growth above the ground. The tuber from which the
stem has developed, and which is therefore in the process of being exhausted,
will be termed the current tuber. The term ‘old tuber’ will only be applied to those
which have been exhausted during a previous season’s growth.

Text-fig. 5.—A longitudinal section of the apex of a ‘dropper’. B, apical
bud; H, scale leaves; V, vascular bundle; L, leaf trace. Growth on the side M
has been greater than on the side N, so that the apical bud is already slightly
to one side. x 17:5.

Text-fig. 6.—A longitudinal section of a young tuber. T, tuber; D, base of
‘dropper’; A, apical bud; E, scale leaves; V, vascular bundles; L, leaf trace.
3g diols).

Text-fig. 7.—A young seedling of D. auriculata. R, root developed from
the radicle; H, root hairs; X, testa; C, cotyledons; A, apical bud; F, foliage
leaves. x 12.

Text-fig. 8—A seedling of D. auriculata with a ‘dropper’. R, root; D,
‘dropper’ arising in the axil of the foliage leaf (A); X, testa; B, apical bud;
O, rosette of foliage leaves. x 5.

Text-fig. 9—A plant of D. auriculata in its second year. Its rosette of
leaves is drawn from below. NT, new tuber; D, ‘dropper’; B, swollen end of
‘dropper’ just above the new tuber; EH, scale leaves; CT, current tuber; R, root;
U, underground stem; F, foliage leaf. x 3:6.

250 VEGETATIVE REPRODUCTION IN DROSERA,

Seale leaves occur at intervals throughout the length of the underground
stem. Elongation of the underground stem or of the ‘dropper’ in the ground
after the formation of the scale leaves, frequently causes them to appear, from a
superficial examination, to point in the opposite direction to that which would be
expected.

Short adventitious roots develop on the underground stem. They are most
numerous immediately above the current tuber, but others occur scattered through-
out its length. Diels (19060) has pointed out in the case of D. erythrorhiza,
that these roots arise at the base of the scale leaves, to which they appear to be
attached. This observation has been confirmed for D. peltata and D. auriculata.
Roots may occur either singly or in twos and threes at the leaf base (Text-fig. 11).
The relatively poor development of the root system may account for the restriction
of these species to rather moist localities.

The ‘dropper’ which is destined to bear the new tuber typically arises in
the axil of a scale leaf immediately above the tuber of the current year, and
penetrates more deeply into the soil. When growing in soil of moderate depth,
the ‘droppers’ usually point vertically downwards; but in the plants which grow
on rock ledges, the ‘dropper’ often becomes coiled and twisted, where its down-
ward progress has been checked by contact with the hard rock. The growing
point of the ‘dropper’ ultimately becomes inverted and swollen to form the new
tuber.

Text-fig. 9 shows a plant of D. auriculata in its second year. The shoot arises
from the apex of the current tuber (CT). It bears a laterally produced ‘dropper’
(D) which is in the process of forming a new tuber (NT) at its extremity. Ata
point immediately above the new tuber, the ‘dropper’ is always considerably
swollen (B, Text-fig. 9). This region was termed the “probulb” by Morrison (1905).
It is connected with the tuber itself only by a narrow neck or connective, but is
usually closely applied to it, as the result of the enlargement of the tissue of the
tuber (cf. Text-fig. 6).

At the end of the second season, the tuber is still fairly small, about 2 mm.
in diameter, but it has been carried considerably deeper into the soil by the
second ‘dropper’, and is thus better able to resist desiccation.

The process of progressive penetration into the soil may go on year by year,
until an appropriate level has been reached. When a certain depth has been
reached, however, the ‘dropper’ as such is no longer formed and, instead, the
new tuber is developed on an extremely short stalk arising from an axillary bud
immediately above the current tuber, i.e., in the same position as the ‘dropper’.
Thus it is situated beside, and at approximately the same level in the ground as,
the current tuber. It is evident from this that the ‘dropper’ is simply an elongated
connective, by means of which the plant is able to adjust its dormant reproductive
shoot to a suitable level below the surface of the ground. The annually produced
tubers are now developed as a rule at this same level.

A longitudinal section through a current and new tuber (Text-fig. 10) shows
the new tuber (NT) on an extremely short connecting stalk (K), which has
developed in the axil of a scale leaf just above the current tuber. Its vascular
tissue arises as a branch from the bundles of the underground stem (U). In
transverse section the underground stem shows a wide cortex, and a ring of
vascular bundles arranged near the centre. Branches from these bundles pass
through the connective and into the new tuber, where they branch again to
form a ring of five or six bundles, which terminate just behind the meristem.

BY JOYCE W. VICKERY.

Text-fig. 10.—A longitudinal section through a current tuber showing the
new tuber developing beside it. CT, current tuber; NT, new tuber; K, connective
branch on which the new tuber is developed; A, scale leaf in whose axil the
branch (K) arose; U, underground stem; EH, scale leaf; R, adventitious root;
V, vascular bundles; G, inner layers of parenchyma of the current tuber which
have contracted away from the outer zone (M). x 12.

Text-fig. 11.—The underground parts of a plant of D. peltata. CT, current
tuber; NT, new tuber; B, scales formed from the remains of old tubers; U,
underground stem; TT, tuber; A, scale leaf in whose axil the tuber (T) arose;
G, ground level; S, aerial stem; R, adventitious roots situated at the base of
the scale leaves. x 1 approx.

Text-fig. 12.—The underground parts of a plant of D. peltata. CT, current
tuber; D, ‘dropper’; NT, new tuber; B, scale formed from the remains of an old
tuber; OT, an old tuber penetrated by the underground stem (U); Y, detritus
formed from the fibrous remains of previous underground stems; R, roots; G,
ground level. x 1 approx.

Text-fig. 13.—An abnormal development of the underground parts of
D. auriculata which had been partially exposed to light. CT, current tuber;
D,, ‘dropper’ bearing a new tuber (NT); R, adventitious roots; D,, a second
‘dropper’ arising on the underground stem near D,; D,, a ‘dropper’ originating as
a branch of D,; A, apical bud of D,; C, apical bud of D,, which is forming a
cluster of foliage leaves; L, level of soil from which D, and D, were projecting ;
U, underground stem; EH, scale leaves; S, rosette of linear scale leaves at ground
level (G). x 1 approx.

Text-fig. 14.—A ‘dropper’ bearing an abnormal tuber of D. auriculata, which
had been partially exposed to light. T, tuber; E, scale leaves; C, enlarged scale
leaves; D, ‘dropper’; F, foliage leaves which have developed on the tuber in
place of scale leaves. x 1 approx.

Text-fig. 15.—An abnormal development of the underground parts of
D. peltata. U, broken underground stem; R, roots; D,, ‘dropper’; D,, a ‘dropper’
which has arisen as a branch of D,; EH, scale leaves; F, foliage leaves which
have developed on the ‘dropper’ in place of scale leaves; L, rosette of linear scale
leaves at ground level; A, aerial stem. x 1 approx.

o1

252 VEGETATIVE REPRODUCTION IN DROSERA,

Two of these are shown in section in Text-figure 10, at V. The connection of
the bundles of the current tuber (CT) with the underground stem of the previous
year is not shown in this section, since they curve upwards in a plane more
or less at right angles to the direction of section.

As the new tuber increases in size the food materials in the current tuber
become exhausted, and the inner tissue (G, Text-fig. 10) shrinks away from the
outer layers (M). The outer zone consists of the epidermis and a few rows
(about three to five) of parenchyma cells, which are torn away from the remainder
of the ground tissue. As the new tuber develops, therefore, the current tuber
becomes soft and pulpy. The young tuber presses in the current tuber more
and more, and so comes to fill the space once occupied by it. The current tuber
finally, when all its foods are exhausted, remains as a dark brown or black
concave shell, adhering to one side of the newly formed tuber (B, Text-fig. 11).
This process may go on at the same level for a number of years, the hardened
shells of the old tubers persisting as outer protective coverings. The young
tubers are formed alternately on different sides of the stem, so that a number
of these black shells or scales may be found completely enclosing the current and
new tubers.

Planchon (1848) tentatively compared these structures to the outer scaly
leaves of the onion, while Morrison (1905) considered that they were an exfolia-
tion of annual occurrence from the substance of the ‘bulb’, and apparently did
not realize that they were the exhausted shells of previous tubers. Their presence
may explain why Morrison and others termed the perennating organ a bulb.

When the apex of the connective becomes inverted to form the new tuber,
the growing point is invariably situated on the side of the new tuber against the
current tuber, never towards the outside. This orientation of the apical bud
causes it to be situated immediately beneath the existing underground stem.
After the aerial parts have died down, this stem becomes soft and rotten,
leaving only a small scar at its point of attachment with the newly formed tuber.
When the growing point starts to shoot, the new stem grows directly upwards,
and so penetrates through the rotting tissues of the old underground stem. It
is possible that the soft central core forms a channel of least resistance to the
developing shoot, which would also be protected from abrasive soil particles by
the disintegrating fibrous outer layers. .

It has been suggested by previous writers (see Rendle, 1925) that this dead
material may function as a velamen does, in holding moisture around the under-
ground stem. This has a strongly lignified epidermis, but the cell walls have
large pits on their outer sides. This structure may allow a certain amount of
water to enter the underground stem over its whole surface.

This detritus of the fibrous remains of previous underground stems can be
seen on all of the older plants (Text-fig. 12), when it may be quite thick, and
frequently on those of only two or three years. Sometimes the underground stem
passes through the remains of old tubers (as at OT, Text-fig. 12), which may
appear to belong to it, but can always be easily separated, and show no point
of attachment.

The presence and cause of this detritus was noticed by Morrison (1905) in
his description of the Western Australian species. It is probable, however, that
when he speaks of the presence of several “bulbs” arranged one above the other,
he mistook the old remains of previous tubers for functional tubers. The appear-

BY JOYCE W. VICKERY. 253

ance was also observed by Planchon (1848), but with the material at his disposal,
he was not able to satisfy himself as to its origin and significance.

Robertson (1906) has described the ‘droppers’ formed by Tulipa and
Erythronium, which carry the new bulb down to a suitable level in the soil, their
length varying accordingly. If the bulb is already placed too deeply in the soil,
the ‘dropper’ may be directed upwards instead of downwards, so that the new
bulb develops higher than the old.

An upwardly directed ‘dropper’ has not been observed by the writer in any
of the plants of D. peltata or D. auriculata examined. One specimen found, how-
ever (Text-fig. 11), illustrated the way in which tubers may be placed at a higher
Tevel in the ground. A tuber (T) has been formed relatively high up on the
underground stem, in addition to the new tuber (NT) formed at the normal
position beside the current tuber (CT).

There is typically only one ‘dropper’ or one young tuber produced every
season by each plant. In just a few cases, however, out of the several hundred
plants examined, multiplication of the underground stem had occurred to form
two ‘droppers’ (e.g., Text-fig. 13), each able to form a tuber. Much more frequently
the ascending axis branches to give two aerial stems, at about ground level, or
just below the surface. This often occurs if the first shoot is injured. Examples
have also been seen in which the apex of the ‘dropper’ had been injured, and an
axillary bud of the ‘dropper’ had grown out and taken its place. Similarly, even
in uninjured plants the ‘dropper’ occasionally branches (Text-figs. 13, 15).

Owing to the rapid disintegration of the old underground organs, this method
of vegetative multiplication can only definitely be observed when branching has
occurred during the current season. Often, however, plants are found growing
in elusters of two or three, with their underground parts in close contact, and
sometimes even growing up through the same fibrous material, but nowhere
actually joined. It seems certain that in many cases these groups have arisen
from the one plant by branching of the underground parts during’ a previous
season, their connection having subsequently disintegrated.

The relationship between the time of appearance of the new tuber or ‘dropper’
and the vegetative and flowering phases of the plant has been investigated. No
direct correlation could be found. Sometimes the new tuber has reached a
considerable size while the aerial part of the plant is still short and with only
leaf bud. At other times, even flowering plants have only an extremely small
new tuber. The two processes then, of tuber and flower production, seem to go
on independently of one another.

Certain abnormal specimens collected are of interest, in that they show the
marked plasticity of the underground parts, and their reactions when subjected to
unusual stimuli.

Several plants of D. auriculata were found growing on a steeply sloping rock
ledge on which the soil was very shallow. Long ‘droppers’ had been tormed,
and had grown completely out of the soil. These were exhibiting some most
unusual features. Text-figure 13 illustrates a typical example of the lower parts
of one of these plants. The current tuber (CT) is situated only a few milli-
metres below the surface of the soil. A young tuber (NT) is already being formed
on the end of a short ‘dropper’, and in addition a second ‘dropper’ has arisen from
a bud on the underground stem close to the first, and has formed a long stolon.
This in turn has given rise to a third ‘dropper’. These last two ‘droppers’ were

254 VEGETATIVE REPRODUCTION IN DROSERA,

free from the soil from about the level L, and were quite green in colour, in
distinction from the usual white appearance. Their scale leaves had become
unusually large, and were quite green. The most extraordinary feature, however,
is shown at C, where the whole apical shoot of the ‘dropper’ has been transformed
into a cluster of small foliage leaves. Every gradation between a normal foliage
leaf with glands, and scale leaves can be seen. The growing point has mean-
while become inverted, as happens during tuber formation. The tuber has thus
been replaced by a small plant, which, however, at this time, had no connection
with the ground. The fact that several ‘droppers’ were produced by many of the
plants, indicates that failure to reach a suitable substratum on the part of the
first ‘dropper’ may lead to the development of other ‘droppers’.

An intermediate case between these clusters of leaves, and the normal tuber,
is shown in Text-figure 14. Here the tuber has been formed, but the scale leaves
on it have become much enlarged, and two of them have developed into foliage
leaves. This transitional stage demonstrates the plasticity of these modified leaves
normally present on the ‘dropper’ and tuber, and establishes the fact that their
scale-like form is the result of their subterranean environment.

Text-figure 15 illustrates a specimen of D. peltata whose behaviour has been
similar to that described above in plants of D. auriculata. At ground level a
rosette of elongated scale leaves can be seen. The ‘dropper’ has branched, and
both ‘droppers’ have developed several foliage leaves with glandular hairs in
place of the usual scale leaves. The underground stem had previously been
injured just below ground level, and only a very small length of stem marks
its former position.

Ecological.

Many plants were found which appeared to have previously reached a
permanent level, having the remains of old tubers about them, and yet had sent
down a ‘dropper’ to a still lower level during the current season. This would
take place in cases where the soil had been somewhat washed away from the
surface, but it also occurred in places where no sign of such a reduction of level
could be detected. The tubers, also, of some of these plants were already at a
considerable depth.

The relation between the depth of the tuber and the habitat, especially the
type of soil in which the plants occur, has been investigated. Measurements of
the underground parts of 263 plants of D. peltata have been made. Only those
plants were measured which were considered to have reached a permanent level
in the soil on account of the presence of old tubers, or of a newly formed tuber
at the same depth as the current tuber.

The results are represented graphically in Text-figure 16. The numbers of
plants from five different localities have been plotted against the depths at which
their tubers occurred below the surface of the ground. The measurements were
made of the length of the underground stem from the ground level to the top
of the tuber. The diameter of the tuber was not included in the measurement,
in order to minimize error due to the size of the tuber.

The absolute heights of the curves are unimportant, since the measure-
ments for the various localities were made on different numbers of plants. Only
the form of the curve and the position of its maximum point is of significance.
The two small secondary maxima in the curves B and E are not thought to be of
significance, but due to error on account of the small numbers used.

BY JOYCE W. VICKERY. 255

Depth tn cm.

Text-fig. 16.—Graph showing variation in depth of tubers of D. peltata
collected from five different localities. A, from Church Point, moist situation,
heavy sandy soil; B, from Waterfall, rather dry situation on top of hill, light
sandy soil; C, from Pennant Hills, moist situation, light sandy soil; D, from
Pennant Hills, very wet situation, heavy sandy soil; H, from Pennant Hills,
rather dry situation on top of hill, clay soil.

Three curves, B, C, and H, have their maximum point at 5 cm. depth in the
soil, the curve A at 4 cm., and the curve D at 3 cm. This result corresponds in
a general way with the moisture content of the soil that would be expected in
the respective localities. The curves B, C, and E were given by plants collected
from areas which would soon become dry in the hot months. The curve A was
given by plants growing near a Swamp on the flat top of a hill, where the soil
would tend to retain its moisture much longer. The curve D is of interest in
that its maximum point occurs at 3 cm. depth, since these plants were collected
in a particularly moist locality which would probably not be subjected to drought
during the summer months. Unfortunately this curve is based on measurements
of only 20 plants, so that too much reliance cannot be placed on it.

These curves indicate therefore that the depth at which the tubers occur is
influenced by habitat. The moisture relations of the soil are apparently of more
significance than the soil texture, since similar curves (C and E) were obtained
from plants on clay and sandy soils respectively.

The average depth of the tuber for the total number of plants examined was
found to be at about 5 cm, below the surface of the ground. In the following
table the numbers of plants which had their tubers at each depth from 2 to 10 cm,
are given:

256 VEGETATIVE REPRODUCTION IN DROSERA,

4

14 51 91

Depth in ecm. Sea Layette ap tia re | 2 | 3
| |
No. of plants | |
}

|

pa]
| |
| |
| |
| |

ADVENTITIOUS Bubs.
i. Epiphyllous Buds.

Accessory methods of vegetative reproduction have been described in various
species of Drosera. Goebel (1918) has described a peculiar method of reproduc-
tion in D. pygmaea. This species produces brood buds which represent modified
parts of the leaf. Proliferation of the inflorescence may also occur. Dixon (1901)
recorded the production of adventitious buds borne in the angles between the
pedicels and the main axis of the inflorescence. Abnormal flowers, sometimes
resembling vegetative shoots, have been described by Planchon (1848), Fernald
(1905) and Leavitt (1905). Levine (1916) showed that sometimes these shoots
could give rise to new plants.

Epiphyllous buds in Drosera were probably first recorded by Naudin (1840)
for D. intermedia. His tentative conclusion that the epidermis was ruptured, and
hence that the buds originated endogenously, does not agree with the findings of
later writers.

Epiphyllous shoots were recorded in D. capensis by Winkler (1903), and in
D. longifolia by Kirschleger (1855). Nitschke (1860) described similar vegetative
budding in D. rotundifolia, and this was recorded in the same species by Graves
(1897), Grout (1898), Dixon (1901), Robinson (1909), and Dennis (1926).

Leavitt (1899, 1901) propagated plants of D. binata, D. dichotoma and D. fili-
formis from adventitious buds on cut leaves. Buds appeared after about three
weeks on leaves placed on sphagnum. Ames (1899) commented on the possi-
bility of propagating certain species of Drosera from old leaves cut from mature
plants. He found that in D. binata, D. filiformis, and D. intermedia americana,
mature leaves which fell on the sand in which the plants were growing produced
adventitious buds. Goebel (1908) also obtained adventitious buds on part of a
leaf of D. binata kept on a moist substratum.

Salisbury (1915) again recorded the occurrence of adventitious buds on the
leaves of D. rotundifolia and D. intermedia on plants which had been kept in a
greenhouse. He noted that the age of the parent leaf on which they occur may
vary.

Other references to adventitious buds in Drosera are made by Godron (1878),
Geisenheyer (1898), and Heinricher (1902).

As far as is known to the writer, the origin and very early stages of develop-
ment of these epiphyllous buds in Drosera have not previously been described.

Epiphyllous buds were first observed by the present author on D. peltata in
November, 1931, when a few plants were found on which several of the foliage
leaves had developed shoots on their upper or ventral surfaces. Some of these
were miniature aerial shoots with a short axis up to 5 mm. in length, bearing
small orbicular foliage leaves. Others appeared as white stolons growing vertically
downwards, suggesting at once the ‘droppers’ of the ordinary plant. Some of
the leaves bore more than one shoot. The parent laminae on which the shoots
occurred were all withered at the time when they were collected. Just previous
to their discovery, there had been a heavy rainfall, and the winter had been a

BY JOYCE W. VICKERY. 257

wet one. Similar shoots were found a few days later on plants growing on
sheltered mossy rocks on the side of a hill, which were kept constantly saturated
throughout the winter by water draining from the ridge above.

Subsequently it was found that these buds could be readily produced under
laboratory conditions, either on leaves detached from the plant, or on uninjured
plants, and thus a constant supply of material for histological examination was
easily procurable.

Epiphyllous buds have been induced in the laboratory on leaves of D. peltata,
D. auriculata, D. spathulata Labill., D. Arcturi Hook., and D. binata Labill. The
last three species have buds resembling those already described by earlier writers.
D. peltata and D. auriculata, however, are unlike those previously recorded in
that sooner or later a ‘dropper’ is developed which bears the young tuber, and
establishes the plant in the ground.

The following description of adventitious buds is given from a detailed
examination of only D. peltata and D. auriculata, but it has been observed that the
buds of D. spathulata, D. Arcturi and D. binata have a similar origin and early
development.

Morphology.

It will be convenient to describe here the typical arrangement of the glands
on the laminae. This is found to be practically the same in both D. peltaia and
D. auriculata. The petiole is attached to the lamina on the dorsal surface at
approximately the centre of the leaf in the adult form of foliage. When the lower
leaves are arranged in a rosette, however, their petioles are attached to the
margins of the leaves and the laminae are more orbicular or semi-orbicular in
shape. A similar shape, usually orbicular, is observed in the leaves of seedlings
or young epiphyllous plants (Text-figs. 4 and 9).

In the adult foliage the whole margin is bordered by long glandular hairs.
Two rows of long stalked glands, an outer and an inner, can be made out at
the margin (Text-fig. 17). Approaching the centre, these are succeeded by a
third row of much shorter glands, whose stalks, however, are rather longer than
the remainder of the glands which occupy the central portion of the leaf and
have only very short stalks. At the two angles of the adult leaf, a number of
the long glandular hairs are united for part of their length to form the ‘auricles’.
These are particularly conspicuous in D. auriculata (Text-fig. 21).

Detached leaves kept in the laboratory on wet sand, or floating in water in
Petri dishes, were examined at frequent intervals, in order to detect the first
outward signs of bud formation. It was found that the bud arises as a swelling
on the upper surface and appears to surround one of the glands completely. In
Text-figure 17 a short gland can be seen emerging from the top of each of the
two very young buds, so that they form swollen enlarged bases to these glands.
At a slightly later stage (Text-figs. 18, 19) the gland is pushed to one side, while
the bud develops a conspicuous apex and continues its growth.

Most frequently this occurs on one of the short glands situated towards the
central portion of the lamina (Text-fig. 18), but occasionally one of the glands
belonging to the inner or even the outer row of marginal glands gives rise to a
bud (Text-fig. 19). In every case, bud formation is associated with a glandular
hair, and consequently shoots always appear on the upper surface.

Only one example has been seen in which the bud appeared, from a super-
neial examination, to arise on the lower side of the leaf (B, Text-fig. 20). This

258 VEGETATIVE REPRODUCTION IN DROSERA,

(ON \ f |
oS
CI a
QW ZS.

DAY

Text-fig. 17.—A leaf of D. peltata bearing two very young epiphyllous buds.
B, buds; G, glandular hair at whose base the bud arose. x 10.

Text-fig. 18.—A leaf of D. peltata bearing two young epiphyllous buds.
B, and B,, buds; G, glandular hairs pushed aside by the growth of the buds;
As apical wbuda, 5 w0:

Text-fig. 19.—A rather juvenile shaped leaf of D. peltata bearing a bud
(B) which has developed from the base of a long stalked gland (G) of the
inner marginal row. The central short glands were withered at this stage. x 6.

Text-fig. 20.—A leaf of D. peltata bearing. a bud (B) apparently on the
lower surface. This bud has developed at the base of the fused gland stalks
(G), and has grown towards the lower surface of the leaf. x 9.

Text-fig. 21.—A leaf of D. auriqulata bearing two epiphyllous buds. 8S, a
slender type of bud; B, a thick bud with short protuberances near its apex.
x 8-5.

BY JOYCE W. VICKERY. 259

bud arose from the base of the fused gland stalks of the ‘auricle’. The gland
stalk is always ultimately pushed aside by the developing bud, and in this case
it was pushed towards the centre, thus causing the bud to appear on the lower
surface.

The buds are sometimes very slender from their earliest appearance (Text-
fig. 18, and at S, Text-fig. 21); at other times they are quite thick with small
protuberances probably representing leaf rudiments (B, Text-fig. 21). Develop-
ment is similar in each case, and the forms cannot be correlated with the types
of buds to be described later.

More than one bud frequently occurs on the lamina. As many as four shoots
have been observed to develop and attain considerable size on the one leaf. An
examination of transverse sections of leaves which have produced one or more
visible buds, has led to the conclusion that any glandular hair, the cells of whose
stalk are in a healthy condition, is potentially capable of giving rise to an
adventitious bud. It is only on healthy leaves, therefore, that buds can be
produced. The withered appearance of the lamina bearing well-developed buds,
that has often been observed, is due to death subsequent to the formation of the
buds. Transverse sections of the leaf often show that numerous gland stalks
have been stimulated, and show early stages of division to form a meristem. It
is probable that one or more meristems, slightly in advance of the remainder,
or with a better food supply, become dominant and inhibit the development of
other meristems.

Two types of buds may occur on the laminae in both species. These cannot
be distinguished until they are about 1 to 2 mm. in length, when they assume a
distinct appearance and habit of growth. The first type resembles in miniature
the aerial shoot of the parent, and is, in all essentials, during the early stages
of its development, similar to those figured by Naudin (1840) and others (Text-
fig. 22). It has an ascending axis on which the small foliage leaves are borne.
These are more or less orbicular, and resemble the leaves first borne by the
seedling. Eventually an axillary bud grows out from the axil of one of these
leaves, and turns downwards to form a ‘dropper’ (Text-fig. 22), similar in all
respects to that which is produced by a seedling. This occurred after one or
two months in buds grown in the laboratory. The ‘dropper’ usually grows over

Text-fig. 22.—An older epiphyllous bud on a leaf of D. peltata. LL, withered
parent lamina on which the bud developed; S, shoot of the epiphyllous bud;
D, ‘dropper’ developed from a bud in the axil of a leaf (A); R, root arising
near the base of the shoot; E, scale leaves; T, tuber. x 2:8.

Text-fig. 23.—An epiphyllous bud which has developed directly into a
‘dropper’ (D). HE, scale leaves; T, tuber. x 4.

Text-fig. 24.—An epiphyllous ‘dropper’ bud (D) which has developed on the
parent leaf (L) while it was still attached to the plant. The ‘dropper’ has
branched at A. Foliage leaves (F) have developed where the ‘dropper’ was
exposed to light. x 1.

Text-fig. 25.—A ‘dropper’ bud (D) which has arisen on the parent leaf (L).
Its apex has become inverted and given rise directly to an aerial shoot (S)
without the formation of a tuber. R, an adventitious root. x 2:5.

Text-fig. 26.—An axillary bud of D. peltata which has formed a ‘dropper’
CD) iairectlya, xolr

Text-fig. 27.—An axillary bud of D. peltata which has formed a leafy shoot
(S). D, a ‘dropper’ which has developed from an axillary bud of this shoot.
R, an adventitious root arising near the base of the shoot; A, leaf scar of the
axillant leaf; O, part of the stem of the parent plant. x 1.

260 VEGETATIVE REPRODUCTION IN DROSEBA,

the side of the parent leaf, or occasionally forces its way through the tissue of
the lamina. The ‘dropper’ may grow one or more centimetres in length and, on
entering the ground, a small tuber is formed (T in Text-fig. 22). When this has
occurred, the adventitious shoot may be considered to have become established
as a separate entity, and is now in the same condition as a seedling plant is
at the end of its first season.

The second type of bud differs from the first in that the apex of the bud
itself becomes a ‘dropper’ at once, and commences to grow straight downwards
towards the soil or substratum, without forming any aerial shoot (Text-figs. 23, 24).
The ‘dropper’ is usually whitish in colour or a very pale green. If it is exposed
to light for some time, as occurs when the parent leaf is some distance from
the ground, the ‘dropper’ turns a stronger green, and the scale leaves sometimes
develop as foliage leaves (Text-fig. 24). Similarly, foliage leaves are sometimes
seen on ‘droppers’ developed on the leafy type of shoot. In some cases the
‘dropper’ has been seen to grow 4 or 5 cm. long (Plate viii, fig. 2), when the
lamina on which the bud arose was situated at some distance above the ground,
and in this case it is usually extremely slender. In a few cases when the ‘dropper’
was unable to reach the substratum, the apex became inverted and gave rise
directly to a small leafy shoot (Text-fig. 25).

Roots were not developed under laboratory conditions until about two months
after the buds had been produced. It was then found that the leafy type of buds
had produced a typical adventitious root with numerous root hairs from their
stems, at a point just above their connection with the parent laminae. If the
root occurred rather higher up on the stem, it was found that it had developed
just at the base of one of the first leaves (R, Text-fig. 22). At the time when
roots are first noticeable, the parent lamina is always very withered. No roots
were ever found on the ‘dropper’ buds, except in the rather unusual case shown
in Text-figure 25, where the ‘dropper’ apex had turned to form a leaf shoot. In
this case a root occurred at about the point where its direction of growth had
changed.

li. Axillary Shoots.

Normal buds present in the axils of the foliage leaves on the aerial stem
have occasionally been observed to give rise to new plants. Under ordinary circum-
stances the dormant axillary bud is extremely small and insignificant, no elaborate
shoot being organized. A ‘dropper’ may either arise directly from the apex of an
axillary bud (Text-fig. 26), or else the axillary bud may produce an ordinary
leafy branch, and a ‘dropper’ may develop from one of the axillary buds of this
branch (Text-fig. 27).

HistToLocy or HPIPHYLLOUS Bubs.

Leaves showing very young stages of adventitious bud formation were
embedded and microtome sections cut. The material was fixed either in acetic
alcohol or in Flemming’s strong fixative, and the majority of the sections were
cut at 10% and mounted serially. Safranin and Delafield’s Hematoxylin were the
chief stains used.

The meristem forming the bud was found to arise from adult cells at the
base of a glandular hair, both in the mesophyll of the leaf, and the epidermal cells
of the upper surface. Sections were cut of fresh leaves just brought into the
laboratory, and of detached leaves which had been kept in water for varying

BY JOYCE W. VICKERY. 261

intervals. It was found that the earliest stage in the appearance of a meristem
could first be detected after about eight days in water.

Text-figure 28 shows a longitudinal section of an adult gland prior to any
meristem formation. An adequate description of the gland in D. rotundifolia
has been given by Huie (1896) as well as by previous investigators, and the
glands of D. peltata and D. auriculata have the same structure.

The earliest stage in regeneration that could be definitely detected is shown
in Text-figure 29. A few cells of the mesophyll of the leaf, near the base of the
gland, are seen to have divided at A, and a few cells of the epidermis (B) also
show signs of recent divisions. At this stage the mesophyll and epidermal cells
are highly vacuolated, with only a thin layer of cytoplasm lining the walls.

c)

is
E\
i)

uN

f
Q

igs

i

shelele.

Rls
Be

x

Text-fig. 28.—A longitudinal section through a mature glandular hair (G),
and transverse section through the lamina. V, spiral vessel which passes up
through the stalk of the gland; C, vacuolated celis at the base of the gland.
Glo OF

Text-fig. 29.—A transverse section of a leaf at the base of a gland showing
the first divisions of the cells to form a bud, in the mesophyll (A) and epidermal
cells (B). x 150. ;

Text-fig. 30.—A transverse section of a leaf and longitudinal section through
a gland showing divisions in the mesophyll (A) and epidermal cells (B) at the
base of the gland. The cells are becoming rather more densely cytoplasmic, and
the original cell walls can be distinguished from those recently formed. x 150.

3 noe.
S 8

262 VEGETATIVE REPRODUCTION IN DROSERA,

Text-figure 30 shows an interesting stage in the development of the bud.
The mesophyll cells (A) at the base of the gland now show numerous divisions,
and the epidermal cells (B) have also undergone several divisions. The difference
in thickness between the old cell walls of the mesophyll cells, which have secondary
lamellae, and the primary walls of the newly formed cells is clearly marked at
this stage. It will be observed, therefore, that each mesophyll cell and certain
of the epidermal cells immediately at the base of the gland are forming meristems
within themselves.

At the stages shown in Text-figures 28 and 29 the intercellular spaces are
large and conspicuous throughout the mesophyll. At the stage shown in Text-
figure 30 they are becoming quite small and inconspicuous in the region of the
meristem. At this stage the cells of the developing meristem are still fairly
highly vacuolated, though certain of the epidermal cells are becoming rather more
densely cytoplasmic.

Ata still later stage a slight protuberance of the leaf surface becomes evident
at the base of the gland (Text-fig. 31), especially on one side (P), so that the
gland stalk is being very slightly pushed to one side. The old walls of the
original mesophyll cells can still be clearly detected in some places (M), but
their complete original outlines cannot now be traced. The exact fate of these cell
walls is uncertain.

Text-figure 32 shows a still later condition. At this stage the bud could be
detected as a small protuberance on the leaf, by means of a hand lens. The stalk
of the gland (G) is situated near the top of the bud, but now definitely over to
one side of it. The meristematic activity has become concentrated on one side
of the gland stalk (M), and this now grows forward carrying the gland with it
for a short time (cf. Text-figs. 18 and 19).

A tangential section of a well developed bud (Text-fig. 33) shows the gland
still attached some distance up the stalk of the bud. It is usually difficult to
obtain a section through an older bud which is median through its meristematic
apex, and also includes the stalk and head of the gland at whose base it arose.
This is because the gland is frequently situated at the side of the growing bud
and at right angles to the plane in which the first leaf is developed. A median
longitudinal section through a similar bud shows that the meristematic apex is
now clearly differentiated, and is giving rise to leaf primordia (Text-fig. 34).

Dixon (1901) and Salisbury (1915) state that the epiphyllous bud is in
vascular continuity with the parent leaf. On the other hand, Winkler (1903)
and Robinson (1909) state that these adventitious buds are not connected with
the vascular system of the parent plant.

A single spiral vessel passes down the stalk of the glandular hair. This
connects the group of tracheides situated in the head of the gland with the
vascular tissue of the leaf, and thus supplies the glandular tissue with water.
This vessel is invariably in connection with one of the vascular bundles of the
leaf, though the ultimate ramifications of the veins may be sometimes too fine
to trace to each gland if the whole leaf be examined externally. Even in trans-
verse section of the leaf, since the vein may be almost parallel to the direction
of section, it is occasionally difficult to demonstrate the actual connection unless
serial sections are carefully examined. The presence of this vessel makes it
apparent that the bud is provided with a rudimentary vascular tissue from the
beginning of its development. Vascular tissue is subsequently differentiated

BY JOYCE W. VICKERY. 263

behind the growing apex in the usual fashion. At V in Text-figure 34 the central
cells behind the meristem are becoming elongated to form conducting cells. That
the vascular system of the bud is continuous with that of the parent lamina is
also shown in Text-figures 32 and 33.

SR,

oie =
‘

a 2,
eas)
P

Stott
oa

fi
<s

2 SST ATER

(Pees

Recep Aarne

Text-fig. 31.—A later stage in the development of a meristem. Outlines of
the original cell walls can be seen, as at P, and M, but the identity of the
original cells is becoming lost. The cells are now fairly densely cytoplasmic. The
upper surface of the leaf is protruding slightly, as at P. x 150.

Text-fig. 32.—A longitudinal section through a young epiphyllous bud. The
growing point (M) is now situated at one side of the gland stalk (G). V,
vascular tissue being formed behind the meristem; L, parent lamina. x 150.

Text-fig. 33.—A tangential longitudinal section through a bud (B) showing
the glandular hair (G) still attached at some distance up its stem. The vascular
tissue (V) of the bud is connected with that of the parent leaf (L). Starch
grains (S) are present in the cortical cells of the bud, and in the mesophyll
cells of the parent lamina near the bud. x 60.

Text-fig. 34.—A median longitudinal section through a bud. M, apical
meristem; F, first leaf of bud; V, cells behind the growing point differentiating
to form vascular tissue. The vascular tissue of the bud is in continuity with
that of the parent leaf (L). x 123.

264 VEGETATIVE REPRODUCTION IN DROSERA,

Salisbury (1915) stated that the buds that he examined were always situated
close to the main vein of the leaf, with which their vascular tissue was at first
connected. In the present investigation, however, it has been found that, in
D. peltata and D. auriculata, the buds may be connected also with any of the
minor veins of the leaf.

The problem of regeneration of plant cells is one of considerable interest.
Several authors, e.g., Goebel (1903), include in the term regeneration those cases
in which new organs arise from the development of latent root and shoot
primordia. In most cuttings the shoot will arise from an axillary bud if one
is present. To this class also would belong those plants in which new organs
arise from the cambium in cuttings, since the primordia initiating their growth
definitely arise from previously existing meristematic cells. In Bryophyllum
calycinum, Naylor (1932) has shown that dormant primordia occur in the notches
of the leaf throughout its life, and that regeneration occurs from these primordia
when the leaf is detached from the plant.

In many plants also, adventitious shoots may occur from the ends of petioles
of leaf cuttings, and propagation may be carried out in this manner. A general
account of the literature on the production of buds on roots and leaves is given
by Holm (1925). The term regeneration may be more strictly applied to that
class of phenomena in which the development of new organs is initiated by cells
which have previously undergone vacuolation, and hence have reached what
may be termed the adult condition. This type of regeneration is less common
in the higher plants. In the propagation of Begonia, however, shoots may arise
near the cut veins from the callus which is formed over the wound, or, in certain
cases, they are developed by the division of a group of epidermal cells, as recorded
by Hansen (1881).

In the discussion on the origin of the adventitious buds in Drosera it has
been shown that the first divisions occur in cells which are in a highly vacuolated
condition, similar to a mature non-dividing parenchyma cell, and that the cyto-
plasm becomes progressively more dense during subsequent divisions. Drosera,
therefore, exhibits a similar type of regenerative phenomena to that shown by
Begonia. There is no sign of any dormant meristematic cells in the leaf which
could give rise to shoots, as there are in the leaf of Bryophyllum. The cells which
divide first are mesophyll and epidermal cells and are in no way connected with
a cambium.

PHYSIOLOGY oF EPIPHYLLOUS BUDS.

The food materials stored in the leaves of Drosera were investigated with the
view of detecting any differences which might occur between ordinary fresh
leaves and leaves which bore adventitious buds.

All leaves which had been freshly collected, or had been kept in a fairly light
position in the laboratory, gave a strong starch reaction with iodine. This starch
may be present in two forms. Granules can be seen within the chloroplast, often
two or three grains occupying almost the whole volume of the chloroplast. Chloro-
plasts occur in the epidermal cells of the leaf in D. peltata and D. auriculata, as
in other species of Drosera (Solereder, 1908), though they are less plentiful there
than in the cells of the mesophyll.

Starch also occurs as free grains. This is the form in which it is stored in
the tuber, and it is usually plentiful in the ‘dropper’, but it is not very abundant
in the fresh leaf, A little may often be seen in cells close to the vascular bundles.

BY JOYCE W. VICKERY. 265
When an adventitious bud arises, however, the cells in its vicinity become a
centre of carbohydrate concentration, and numerous grains of starch are found
in them (Text-fig. 32). As development of the bud proceeds, this starch is
apparently transterred to its tissues, where it is stored in small grains in the
cortex (Text-fig. 33). This abundance of starch in the developing bud was also
noted by Salisbury (1915). No starch reaction was given by leaves which had
been kept in darkness for 24 hours or longer.

A slight tannin reaction was shown by the contents of the cells of the gland
stalk and head when tested with ferrie chloride.

Fat was found to be very plentiful in almost all of the leaves examined. It
is found in the cells of the gland heads, in the epidermal cells of their stalks, and
throughout the epidermis of the lamina. Occasional cells of the mesophyll also
contain it, especially those near the veins. In the epidermal cells of the long
stalks of the marginal glands, it occurs in large globules up to 20u in diameter.
In other cells it may be in globules varying from a similar size to such small
particles that, under relatively low magnification, it appears to be diffused through-
out the cytoplasm.

This fat can be readily demonstrated in fresh sections with osmic acid, with
which it immediately stains black. It is readily soluble in alcohol, as well as in
the usual fat solvents such as chloroform and ether. It appears to be in two
forms in the leaf. That in the epidermis of the smaller glands is soluble in
chrom-acetic fixative, so that leaves fixed in Flemming’s strong fixative (osmic
acid, chromic acid and acetic acid) did not show fat in the epidermis when
subsequently sectioned, although they appeared dark in colour. The large globules
of fat in the long stalks of the marginal glands did not dissolve in Flemming’s
fixative. The fat gives a blue colour with Nile Blue Sulphate, suggesting that it
is of the nature of a fatty acid. A rather lighter fat reaction was given by plants
which had been kept in the laboratory for more than nine months, and whose
present shoot had arisen from the tuber while in a covered glass dish. This may
have been due to the effect on their metabolism of the unnatural conditions in
which they were growing, or it may possibly be associated with their inability to
capture and digest insects. Darwin (1878) showed that plants of D. rotundifolia
which were allowed to capture insects, or were fed on small pieces of meat, became
more vigorous in their growth than plants which were not supplied with organic
material.

Certain other bodies appear in some cells of the leaves and other parts of the
plant body, whose nature and origin could not be definitely established. These
appear as spherical globules, varying in size up to about 10u. They are usually
single, but sometimes in clusters as though they were in contact, suggesting that
they are in at least a semi-solid state. The larger bodies are black in colour, or
an olive-green in thin microtome sections. Many of the smaller ones appear
reddish-brown, but they are thought to be all of a similar nature.

These bodies do not change colour with osmic acid, and are insoluble in
alcohol, chloroform, ether, or dilute or concentrated hydrochloric acid. On account
of their strong natural colour, few tests involving colour reactions could be used.
With Nile Blue Sulphate they stained blue in thin section. it is thought that
they are probably hydrocarbons formed during the metabolism of the fats,
especially since they occur chiefly in the epidermis where the fats are most
abundant. The possibility that they represent decomposition products is suggested

266 VEGETATIVE REPRODUCTION IN DROSERA,

by the fact that they were found much more abundantly in leaves which had
become withered and were being attacked by saphrophytic fungi under the humid
conditions prevailing in the laboratory experiments. They are not invariably
present, as sections of some leaves did not show any of these bodies.

It has been observed by several previous writers (Grout, 1898; Graves, 1897;
and Salisbury, 1915) on the epiphyllous buds of Drosera, that their production
appears to be associated with very moist conditions. Dixon (1901) asserts, on the
other hand, that adventitious buds appear only when the plant is allowed to dry
out. He states that they were produced within two months on plants, growing
on sphagnum under a bell-glass, which were allowed to dry out. It seems to the
present writer, however, as far as can be judged from his brief description, that
the atmosphere under the bell-glass would become extremely humid. Under such
conditions, it has been found in the present investigation, buds may be detected
after 14 days on some leaves.

A number of experiments have been carried out on D. peltata in an endeavour
to ascertain the nature of the stimuli which cause or permit the regeneration of
tissue, and hence the development of the buds. The results of these are sum-
marized below.

It was found early in the present investigation that abundant moisture
favoured the production of epiphyllous buds. In the field they have only been
found in extremely wet situations. They are produced readily on the leaves of
plants kept in glass dishes with bell-covers in the laboratory, though their appear-
ance under these conditions is rather slower than when the leaves are detached
from the plant. Plate viii, figures 1 and 2, shows buds on plants kept under these
conditions. Numerous buds appear, after two or three weeks, on detached leaves
(laminae and petioles) placed on wet sand, or floating on water. In the latter
case, often 50% or more of the leaves could be induced to produce buds.

When stem cuttings including at least one node and leaf were placed under
moist conditions, usually an axillary bud grew out and formed an aerial shoot.
Occasionally an epiphyllous bud was produced. In Begonia and Bryophyllum,
epiphyllous buds are not usually produced if an axillary bud is present. It is
rather surprising, therefore, that in D. peltata epiphyllous buds are so readily
produced on complete plants.

Leaves which had been injured, e.g., by the removal of a few gland heads,
were still able to produce buds, but these always arose from an uninjured gland,
away from the injured part.

Ames (1899) stated that a low temperature was favourable to the production
of buds. In D. peltata it has been found that variation of temperature does not
appear to have any noticeable effect within the limits experienced in the labora-
tory, i.e., between about 12° and 26° C. No controlled experiments were carried
out to test this factor, but buds occurred equally well during the early summer
and the middle of the winter.

The buds are produced most readily in fairly bright light. Leaves placed in
darkness or in deep shade did not show any signs of regeneration.

Leaves fed with minute pieces of meat became contaminated with bacteria
and had to be discarded. It is unlikely, however, that the metabolism associated
with insect digestion is necessary for bud production, since young leaves which
opened under cover in the laboratory frequently formed buds. Leaves fed by
placing in a 2% solution of glucose produced no buds, but turned a deep red
colour, and their glands all became bent: towards the centre of the leaf.

BY JOYCE W. VICKERY. 267

Buds most frequently arise on leaves of the typical adult form, but occasionally
they are noticed on the more orbicular shaped leaf characteristic of the rosette
leaves. In order to test whether the older or younger leaves produced buds more
readily, leaves were detached from 40 plants and placed on wet sand in the order
in which they occurred on the plant, all living leaves being included. It was
found that the highest proportion of buds occurred in the upper third of the
plant. The middle leaves produced fewer buds, but were in turn more productive
than those belonging to the lower third of the plant. This result can be directly
correlated with the healthy appearance of the glands on the leaves, since the
glands of the lowest leaves are often slightly withered when the leaf is still living.
On the other hand, young leaves near the apex, which were only just unfolding,
frequently produced buds.

The only factor. therefore, which can be definitely stated to be essential for
bud production is that of abundant moisture. Very humid conditions, with conse-
quent reduction of transpiration, favour the initiation of a meristem.

SUMMARY.

D. peltata and D. auriculata are winter herbs common in the Sydney district.
They perennate by means of subterranean tubers which occur at the base of
underground stems.

The annual development of the tubers on the plant is described, commencing
from the germination of the seed. A ‘dropper’ arises from an axillary bud of
the seedling, and penetrates for a short distance into the ground. A tuber develops
at its apex by an inversion of the growing point and an enlargement of the cortical
tissues. The next season’s shoot arises from this tuber. A ‘dropper’ may be
formed each season for several years, until the tuber is situated several centi-
metres below the surface of the ground. After this, the new tuber is formed at
the same level as the current tuber each season.

The shoot developing from the tuber at the beginning of the season penetrates
through the remains of the old underground stem, which persists as a loose
fibrous sheath around it. The ‘dropper’ and the underground stem bear small
colourless scale leaves. The underground stem bears a number of short adven-
titious roots, densely covered with root hairs.

Several abnormal specimens are described which illustrate the plasticity of
the underground parts. If exposed to light, foliage leaves may develop on the
‘dropper’ in place of scale leaves.

Branching of the ‘dropper’ may occur, resulting in the formation of two
tubers. Sometimes a second tuber may be formed from a bud on the under-
ground stem some distance above the current tuber.

The relation between the depth of the tuber and its habitat is considered. A
variation diagram is given for several localities, which illustrates the effect of
habitat on the depth at which the tuber is placed when a permanent level has
been reached by means of the ‘droppers’.

Accessory methods of vegetative reproduction are described. Axillary buds on
the aerial stem may occasionally give rise to a ‘dropper’, or to a leafy shoot on
which ultimately a ‘dropper’ develops from an axillary bud. An adventitious root
develops at the base of the stem of the leafy type of bud after about two months.

The epiphyllous buds originate by the division of adult vacuolated cells
situated immediately at the base of a glandular hair. These become meristematic
and the bud grows up, pushing the gland to one side. The vascular tissue of

268 VEGETATIVE REPRODUCTION IN DROSERA,

the bud is in continuity with that of the parent lamina. Starch accumulates in
the leaf in the vicinity of the bud, and passes into the cortical cells of the bud.

Epiphyllous buds arise either on detached or undetached leaves when placed
in very moist conditions.

In conclusion, I wish to express my thanks to Professor T. G. B. Osborn, of
the Department of Botany, Sydney University, for suggestions and kindly criticism
throughout the course of this investigation.

Literature Cited.

*AMES, O., 1899.—An Hasy Method of Propagating Drosera filiformis. Rhodora, i, p. 172.

BENTHAM, G., 1864.—Flora Australiensis, ii, pp. 453-470. London.

DARWIN, F., 1880.—Experiments on the Nutrition of Drosera rotundifolia. Journ. Linn.
Soc. London, xvii, Bot.

DENNIS, M., 1926.—Le Bouterage Epiphylle des Drosera. La Feuwille des Naturalistes,
N.S., No. 27, pp. 67-70. Mai.

Diets, L., 1906a.—Die Vegetation der Erde. vii. Die Pflanzenwelt von West-Australien
Stidlich des Wendekreises. Leipzig.

, 1906b.—Blattrhizoiden bei Drosera. Ber. Deutschen Bot. Ges., 34, pp. 189-191.

Dixon, H. H., 1901.—Adventitious Buds on Drosera rotundifolia. Notes from Bot.
School, Trinity Coll., Dublin, pp. 144-145.

Drupbg, C., 1891.—In Engler and Prantl, Die Naturlichen Pflanzenfamilien. Teil iii, 2,
pp. 261-272. Leipzig.

Ewart, A. J., 1930.—Flora of Victoria, p. 553. Melbourne Univ. Press.

*FERNALD, M. L., 1905.—A Peculiar Variety of Drosera rotundifolia. Rhodora, 7, pp. 8-9.

*GBISENHEYER, L., 1898.—Knospenbildung auf Blattern. Deutsche Botan. Monatschrift,
xvi, pp. 133-134.

*GoDRON, D. A., 1878.—Etudes sur les proliferations. Mem. Acad. Stanislas, 1877. Ref.
in Bull. Soc. Bot. France, 1878.

GOEBEL, K., 1903.—Regeneration in Plants. Bull. Torr. Bot. Club, 30, No. 4, pp. 197-205.

, 1908.—Hinleitung in die Experimentelle Morphologie der Pflanzen, p. 196.
Leipzig and Berlin.

, 1913.—Organographie der Pflanzen, pp. 96-97. Jena.

GRAvEs, J. A., 1897.—Notes on Drosera. Plant World, i, p. 28.

Grout, A. J., 1898.—Adventitious Buds on Leaves of Drosera rotundifolia. Amer. Nat.,
B74) Joh Teale

HANSEN, A., 1881.—Vergleichende Untersuchungen tiber Adventivbildungen bei den
Pflanzen. Abh. Senckenberg. naturf. Ges., 12, pp. 147-198.

*HEINRICHER, E., 1902.—Zur Kenntnis von Drosera. Zeitschrift des Ferdinandeums, 3
Folge, 46 H., pp. 1-30.

Houm, T., 1925.—On the Development of Buds upon Roots and Leaves. Ann. Bot., 39,
pp. 867-881. :

Huis, Liny, 1896-7.—Changes in the Cell-organs of Drosera rotundifolia produced by
Feeding with Egg-albumen. Journ. Microscopical Sci., N.S., xxxix, pp. 387-425.

KIRSCHLEGER, M., 1855.—Note sur quelques anomalies végétales. Bull. Soc. Bot. France,

pp. 355-359.
*LEAVITT, R. G., 1899.—Adventitious Plants of Drosera. Rhodora, i, p. 206.
*__________. 1905.—Translocation of Characters in Plants. Rhodora, 7, pp. 13-20, and 21-31.
*_______ 1909.—-Seedlings and Adventitious Plants of Drosera. Torreya, ix, pp. 200-201.

LEVINE, M., 1916.—Further Observations on Chloranthy in Drosera intermedia. Bot.
Gaz., 62, pp. 389-392.

Lusppock, J., 1892.—On Seedlings, p. 517. London.

Morrison, A., 1905.—Note on the Formation of the Bulb in West Australian Species of
Drosera. Trans. Proc. Bot. Soc. Hdinburgh, 22, pp. 419-424.

,1907.—Further Note on Australian Tuberous Droseras. JIbid., pp. 236-237.

NAvuDIN, M., 1840.—Note sur des Bourgeons nés sur une Feuille de Drosera intermedia.
Ann. Sci. Nat., Bot., Seconde Série, t. 14, pp. 15-17.

Naytor, E., 1932.—The Morphology of Regeneration in Bryophyllum calycinuwm. Amer.
Journ. Bot., xix, pp. 32-40.

*NITSCHKE, T., 1860.—Wachsthumverhaltnisse des rundblattrigen Sonnenthaues. Bot.
Zeit., 18, pp. 57-69.

* These papers were not available to the author.

Proa. Linn. Soc, N.S.W., 19338. PLATE VIII.

Drosera peltata.

ts , i
dE a ry
Sea]
‘ : ‘
-
mm

BY JOYCE W. VICKERY. 269

PLANCHON, J. E., 1848.—Sur les Droseraceae. Ann. Sci. Nat., Bot., 3° Série, t. 9, pp.
79-99.

RENDLE, A. B., 1925.—The Classification of Flowering Plants. Vol. ii. Dicotyledons.
Pp. 1938-199. Cambridge University Press.

RROBERTSON, AGNES, 1906.—The ‘Droppers’ of JTulipa and Hrythronium. Ann. Bot., xx,

pp. 429-440.
ROBINSON, WINIFRED, 1909.—Reproduction by Budding in Drosera. Torreya, ix, pp.
89-96.

SALISBURY, E. J., 1915.—On the Occurrence of Vegetative Propagation in Drosera. Ann.
Bot., xxix, pp. 308-310.

SOLEREDER, H., 1908.—Systematic Anatomy of the Dicotyledons. (Trans. by Boodle
and Fritsch.). Vol. i, p. 326. Oxford.

WINKLER, H., 1903.—Uber regenerative Sprossbildung auf den Blattern von Tovenia
asiatica L. Ber. Deutsch. Bot. Ges., 21, pp. 96-107.

DESCRIPTION OF PLATE VIII.

Fig. 1.—Two leaves of D. peltata bearing epiphyllous buds. ‘The leaf on the left
has a leafy shoot on which a ‘dropper’ has developed. The leaf on the right has a leafy
and a ‘dropper’ bud. x 1:5.

Fig. 2.—A plant of D. peltata on which several ‘dropper’ and leafy epiphyllous buds
have been produced. x 1:5.

ON THE DISTRIBUTION, HABITAT AND REPRODUCTIVE HABITS OF
CERTAIN EUROPEAN AND AUSTRALIAN SNAKES AND LIZARDS,
WITH PARTICULAR REGARD TO THEIR ADOPTION OF VIVIPARITY.

By H. CiairE WEEKES, D.Sce., Linnean Macleay Fellow of the Society in Zoology.
(One Text-figure. )

[Read 26th July, 1933.]

Introduction.

The data used for this paper were obtained while collecting lizards and snakes
for an examination of their placentation.

I wish to thank both Professor A. N. Burkitt, Department of Anatomy, and
Professor C. Witherington Stump, Department of Embryology and Histology,
University, Sydney, for reading and criticizing the paper.

I also wish to thank the Royal Society of London for a grant enabling me to
investigate the snakes and lizards in the Pyrenees and the French Alps, and the
Linnean Society in New South Wales for grants for collecting in Tasmania and
south-eastern Australia.

Material.

Investigations were made of the distribution, habitat and reproductive habits
of snakes and lizards and the climatic conditions under which they breed over
the coastal plain, the Great Dividing Range and the inland plain of south-eastern
Australia during five breeding seasons, and in Tasmania over one breeding season,
covering in all a distance of about 3,000 miles, in a north-south direction of about
1,000 miles (Text-fig. 1). Similar investigations were made in the Pyrenees and
the French Alps during the breeding season of 1931.

Viviparity was found in the Australian lizards Lygosoma (Liolepisma)
entrecasteauxi, L. (L.) weekesae, L. (L.) ocellatum, L. (L.) metallicum, L. (L.)
pretiosum, L. (Hinulia) quoyi, L. (Hemiergis) quadridigitatum, L. (Hemiergis)
decresiensis, Tiliqua nigrolutea, Egernia cunninghami, E. striolata, EH. whitei,
EK. kingi, and in the snakes Denisonia superba, D. suta, Notechis scutata and
Pseudechis porphyriacus.

The proportion of viviparous to oviparous species of reptile ‘on the inland
plain and on the Great Dividing Range of south-eastern Australia was extra-
ordinarily high. On the inland plain the commonest lizards seen and collected
were the viviparous Egernias, EL. cunninghami, E. striolata, E. whitei and E. kingi.
It was not uncommon to find small stony hills with their reptilian fauna entirely
restricted to viviparous species. £#. striolata completely monopolized, as far as
lizards were concerned, the hills around Narrabri. Few oviparous species were
seen throughout the entire expedition on the inland plain, although many species
known to be oviparous have been recorded from the localities visited.

BY H. CLAIRE WEEKES. 271

On the Dividing Range the viviparous species were again by far the most
prolific. At Jenolan, 4,000 feet above sea-level, and about 100 miles west of
Sydney, of the twelve species of reptiles collected ten were viviparous. These
were the lizards 2. cunninghami, H. striolata, EH. whitei, L. (L.) entrecasteauzi,
L. (L.) weekesae, L. (Hemiergis) quadridigitatum, L. (Hinulia) quoyi, T'. nigro-
lutea, and the snakes Pseudechis porphyriacus and Notechis scutatus. Only two
specimens of the oviparous lizard L. (L.) quichinoti were collected and about a
dozen oviparous Amphibolurids were seen, whereas it was easy to catch with the
hand as many as forty viviparous specimens in one day.

At Mount Kosciusko, at 5,000 feet above sea-level, the viviparous L. (L.) entre-
casteauxi and L. (H.) quoyi were very prolific, while only one oviparous specimen,
an Amphibolurid, was seen at this height, although they were numerous lower
down on the mountain slopes.

At Mount Wellington, Tasmania, at 4,000 feet above sea-level, L. (L.) pretiosum,
L. (L.) ocellatum and L. (L.) metallicum, all viviparous lizards, occurred in
hundreds, and yet not one oviparous specimen was seen.

It was only on the coastal plain of south-eastern Australia and on the lower
slopes of the Great Dividing Range up to 3,000 feet above sea-level that the number
of oviparous species approached that of the viviparous.

TENTERFIELD

GUY FAWKES y--"7~
NARRABRI a”

NASSiaee
WAGGA @¢-----&
os t

. y
Sra Cc,

MT KOSCIUSKO’

MT WELLINGTON

Broken line indicates routes followed in collecting in south-eastern
Australia and Tasmania.

272 CERTAIN SNAKES AND LIZARDS AND ADOPTION OF VIVIPARITY,

On the lower slopes of the Pyrenees up to about 3,000—4,000 feet above sea-level
the oviparous lizards Lacerta viridis, L. agilis and L. ocellatus were numerous;
above 4,000 feet they were much rarer, while at 5,000 feet, except for an occasional
L. viridis, they were absent; whereas at 5,000 feet, L. vivipara, a viviparous
Lacerta, occurred and was prolific. The same condition prevailed in the French
Alps. At low levels the oviparous species were prolific, whereas at 6,000 feet,
above Chamonix, only L. vivipara was seen.

The scarcity of oviparous lizards on the inland plain of south-eastern Australia
is not surprising when the prevailing climatic conditions are taken into considera-
tion. During the breeding season of the reptiles and the months of pregnancy,
November, December, January and February, the inland plain is usually very
hot and dry, and species with parchment shelled eggs exposed to the danger of
desiccation have supposedly fewer chances of survival than viviparous species with
eggs safely carried within the female.

At Barrington Tops, 4,500-5,500 feet above sea-level, in January, 1925 (mid-
summer), wintry conditions prevailed while the viviparous reptiles were at the
middle of their gestation period; at Mount Wellington, also in midsummer, snow
was still lying on the ground where the viviparous lizards were carrying their
young in advanced stages of development, and at each of the regions of 4,000 feet
and over visited the nights are normally cold. So that cold is the most obvious
factor which would interfere with the successful development of eggs of oviparous
species at these heights and hence with the successful establishment of such species
at such heights.

On the mountain slopes and on the coastal plain of south-eastern Australia,
however, the climatic conditions are most favourable for the establishment of
oviparous species.

It is interesting that no viviparous species of lizard or snake was found or
has been recorded restricted to the inland plain. Each viviparous species occur-
ring on the inland plain also occurs at at least 4,000 feet above sea-level on the
Great Dividing Range.

The absence of any viviparous species restricted to the inland plain is not
surprising when the factors concerned in the change from oviparity to viviparity
are considered. In reptiles adopting viviparity the eggs are retained in early
stages of development for longer and longer periods in the oviducts and gradually,
in the course of evolution, laid in more advanced stages of development surrounded
by a soft shell, which becomes softer and softer at the time of laying as the
time of retention is lengthened. It is difficult, therefore, to conceive how vivi-
parity could have been evolved in places as hot and dry as the inland plain of
eastern Australia is during the breeding season of the reptiles, since the mortality
of the soft-shelled eggs from desiccation would surely be fatal to the survival of
the species. It was at first difficult to understand how, if the adoption of vivi-
parity under hot dry conditions is unlikely, such desert species as Chalcides
sepoides in Egypt could have evolved their viviparity, until it was learnt that in
Egypt at least the reptiles breed in winter and not in the hot summer months.
While a change of breeding season is practicable in regions where there are months
sufficiently mild to allow for breeding activities of the cold-blooded reptiles, it is
not practicable in such regions as south-eastern Australia, where the winter
months are too cold for breeding. It would be particularly interesting if precise

BY II. CLAIRE WEEKES. 273

information were available on the viviparous reptiles of South Africa from the
point of view of the climatic conditions under which they breed.

Conditions on the Great Dividing Range and its slopes and on the coastal
plain of south-eastern Australia would be favourable for the passage through such
soft-shelled stages, since in these regions danger from desiccation would be
slight. It is curious that under these circumstances no viviparous species has
yet been found restricted to either the mountain slopes or the coastal plain. As
on the inland plain, every species which occurs on the mountain slopes or the
eoastal plain also occurs on the mountain tops. Two viviparous species of lizard
are, however, restricted to 4,000-7,000 feet above sea-level. These _ species,
L. (L.) entrecasteauxvi and L. (L.) weekesae, are the only reptiles recorded from
Australia, and two of the three recorded in the world, which have eggs with a
reduced yolk-content and a specialized, folded, glandular area of allantoplacenta-
tion (Harrison and Weekes, 1925; Weekes, 1929). In these two species and in
the third species, the Italian lizard Chalcides tridactylus (Giacomini, 1891), the
reptilian reproductive cycle attains its greatest specialization. Since the speciali-
zation has advanced so much further in these species than in other species, it is
a natural conclusion that they either began to evolve their viviparity earlier than
other species or else have evolved it more rapidly. Therefore it seems possible
that in south-eastern Australia the factors determining the adoption of viviparity
may be either (a) definitely associated with the altitudes to which these lizards
are restricted, or (0) may work most efficiently under conditions prevailing at
such heights.

In other words, is viviparity among the reptiles from south-eastern Australia
an expression of their adaptation to some external factor or factors associated
with high altitudes, or is it caused by some pathological upset or mutation among
the reptiles, for the evolution of which the conditions at high altitudes are most
favourable?

It is inconceivable that mere barometric pressure could directly influence the
adoption of viviparity. It seems more likely that if there is an external factor
associated with high altitudes influencing the adoption of viviparity, that factor
is cold, whether working by directly interfering with the oviparous cycle or by
merely supplying conditions under which some mutation towards viviparity can
work best.

The Geckos are an interesting group from this point of view. Their distribu-
tion is chiefly tropical and subtropical and they are oviparous, laying eggs with
calcareous shells. New Zealand, with its climate approaching that of the British
Isles, is the coldest country in which they occur. The two New Zealand Geckos,
Hoplodactylus pacificus and Nautilus elegans, are viviparous and they are the only
viviparous Geckos so far recorded in the world.

An investigation of lizards around Sydney during the beginning of the breeding
season of 1926 showed that cold does apparently retard the development of the
eggs within the ovary. During the hot spring of 1926 the development of the
eggs within the ovary, their ovulation and appearance within the oviducts took
about two weeks, while during the colder spring of 1927 the completion of the
same processes in the same species in the same localities took nearly four weeks.
It seems conceivable that, since the cold-blooded reptiles depend so much on
external warmth for the completion of the early phases in their reproductive cycle,
during such cold spells as so often occur in the spring at 4,000 feet and over

274 CERTAIN SNAKES AND LIZARDS AND ADOPTION OF VIVIPARITY,

above sea-level on the Great Dividing Range, cold may directly interfere with
the mechanism controlling the laying of the eggs to retard laying and cause
retention.

It is suggested in conclusion that the general observations in this paper show
the desirability of further investigation, both analytical and experimental, into
the occurrence of viviparity among reptiles.

SUMMARY.

1. This paper is concerned with the results of investigation of the distribution,
habitat and reproductive habits of certain lizards and snakes in south-eastern
Australia, the Pyrenees and the French Alps.

2. It was found that the proportion of viviparous to oviparous reptiles on
the Great Dividing Range and the inland plain of south-eastern Australia was
extraordinarily high, and the number of oviparous species only approached the
number of viviparous species on the lower slopes of the Dividing Range and on
the coastal plain.

3. In the Pyrenees and French Alps by far the majority of reptiles found at
high altitudes were viviparous. The oviparous species were practically confined
to the mountain slopes up to about 4,000 feet above sea-level.

thy he 1S suggested that the failure of the oviparous species to establish them-
selves on the inland plain of south-eastern Australia is due to desiccation of their
eggs by the heat.

5. It is suggested that the failure of the oviparous species to establish them-
selves at 4,000 feet and over above sea-level is due to cold interfering with the
development of the eggs in the nest.

6. All viviparous species on the inland plain of south-eastern Australia also
occur at at least 4,000 feet above sea-level on the Great Dividing Range.

7. Two viviparous species of lizard with the most highly specialized allanto-
placenta and reproductive cycle yet recorded for reptiles in Australia, and only
equalled by the lizard Chalcides tridactylus in Italy, are restricted to 4,000-7,000
feet above sea-level.

8. Therefore it is suggested that, in south-eastern Australia at least, the
factors determining the adoption of viviparity may be either (qa) definitely asso-
ciated with the altitudes to which these lizards are restricted or (b) may work
most efficiently under the conditions existing at such altitudes.

9. Cold is suggested as the most likely external factor associated with high
altitudes that may influence, either directly or indirectly, the adoption of viviparity.

Bibliography.
GIACOMINI, 1891.—Matériaux pour l’étude du développement du Seps chalcides. Arch.
Ital. de Biol., xvi, pp. 332-359.
HARRISON and WEEKES, 1925.—On the Occurrence of Placentation in the Scincid Lizard
Lygosoma entrecasteauxvi. Proc. LINN. Soc. N.S.W., 1, pp. 472-486.
WEEKES, 1929.—On Placentation among Reptiles. No. I. Proc. Linn. Soc. N.S.W., liv,
part 2.

A NEW GENUS AND SPECIES OF AUSTRALIAN PROCTOTRYPIDAE.

By ALAN P. Dopp.
(One Text-figure.)

[Read 30th August, 19338.

The family Proctotrypidae is not rich in genera; six are recognized in
Kieffer’s monograph (Das Tierreich, Berlin, 1914), but it is doubtful whether
Phaenoserphus Kieffer and Cryptoserphus Kieffer are sufficiently distinct from
Proctotrypes Latreille (=Serphus Schrank) to warrant generic rank. The Aus-
tralian Acanthoserphus Dodd, 1915, has since been added to the number of genera.

Nine species in this small family have been described from Australia. Hence,
the recognition of a new genus and species is a matter for comment; due credit
should be given to the discoverer of this interesting form, Mr. F. Erasmus Wilson.
The new insect is Braconid-like in general appearance, and possesses outstanding
characters, particularly in regard to the development of the wing venation.

AUSTROSERPHUS, new genus.

9, g.—Head from dorsal aspect transverse; from frontal aspect as long as
its greatest width, sub-triangular; eyes prominent, bare; ocelli rather close together
in centre of vertex; immediately above the antennal insertions the frons is
produced in the form of a broadly convex transverse lamina, its margin lightly
carinate, its surface broadly depressed medially; a median carina separates the
antennal insertions and meets the frontal lamina; maxillary palpi very long,
5-jointed, the first joint short; labial palpi short, 3-jointed; mandibles not large,
the one edentate, its tooth broadly curved, the other bidentate with a broadly
curved inner and small outer tooth. Antennae 13-jointed in both sexes; scape
stout, its inner dorsal edge carinate, its dorsal surface produced forward in a
triangular acuminate process which covers the pedicel and base of the first
flagellar joint; pedicel small; flagellar joints cylindrical, the basal ones long.
Pronotum short, its anterior border margined, its latero-anterior angles sub-
acuminate; scutum long, the parapsidal furrows complete and almost meeting
posteriorly; scutellum large, at base with a large transverse fovea sub-divided
into five smaller foveae by longitudinal carinae, posteriorly with a row of foveae;
metanotum rather strongly depressed on either side of the sub-quadrate median
area, its posterior margin with four short teeth at wide intervals which correspond
to similar teeth on the anterior margin of the propodeum; propodeum distinctly
separated from the metanotum, on one plane, partially rugose. Forewings ample;
with very complete venation; subcostal vein well separated from the costa; stigma
slender, twice as long as wide; radial vein rather short, at right angles to the
stigma; radial cell closed, large, longer than the stigma; first cubital vein
complete and straight to the distal wing margin; vein Cu. + 1A rather near the
posterior margin which it joins beyond the middle; veins M,,. and M;., almost
join proximately, the former running obliquely to the anterior margin, the latter
running straight to the centre of the distal margin; two medio-cubital and two
inter-cubital veins are present. Hindwings with a costal vein only, except for

276 NEW GENUS AND SPECIES OF PROCTOTRYPIDAE,

a false cross-vein which extends for two-thirds the width from the posterior margin.
Legs normal; posterior coxae, femora, and tibiae rather slender, the tibiae with
two short stout apical spurs; tarsal claws strongly curved, simple. Abdominal
petiole in the female stout, as wide as long; in the male very long and slender,
fully one-half as long as body of abdomen. Body of abdomen somewhat compressed
dorso-ventrally; from lateral aspect abruptly declivous apically in the male;
composed of four segments in the female, five in the male, of which segment 2
(first body segment) occupies more than one-half the surface; in the female
produced in a very slender, cylindrical, straight oviduct, which is longer than
the main abdomen.

Genotype, A. albofasciatus, new species.

A fine genus, distinguished by the very complete venation, the produced
acuminate scape, and the long abdominal petiole in the male. Acanthoserphus
Dodd possesses the produced scape and the complete venation except that M,,,
and Cu, are false and not true veins, but the abdominal petiole is short in both
sexes, the pronotal angles are toothed or spined, and the metanotum bears a long
spine.

AUSTROSERPHUS ALBOFASCIATUS, ND. SD.
9.—Length, to apex of oviduct, 8:50 mm.; to base of oviduct, 5-75 mm.
Shining black, including the antennae and legs, except as follows: ninth and tenth
antennal joints, basal third of intermediate and posterior tibiae, and third and
fourth joints of intermediate and posterior tarsi, silvery-white; similar portions
of the anterior tibiae and tarsi, and the palpi, whitish; oviduct deep ferruginous.

fa, +A
mez
Forewing of Austroserphus albofasciatus Dodd.

Antennae not much shorter than the body; funicle 1 as long as the produced
scape, 2-10 gradually shortening, 10 a little more than one-half as long as 1;
apical joint almost as long as 1, very broadly rounded at apex. Body impunctate,
shining, except the propodeum and metapleurae; with a scattered fine pubescence,
the face below the antennal insertion densely shortly pubescent. Scutum with a
fine median carina on posterior half; propodeum smooth for basal third and
with smooth areas, except for fine surface sculpture, on either side of the median
line posteriorly, the rest punctate to rugose-punctate; posterior two-thirds of
propodeum with strong lateral carinae which are obliquely angled at one-half
their length, and with two or three median carinae, the lateral margins with

BY A. P. DODD. 277

fine long hairs, the posterior margin convex and carinate; metapleurae densely
rugose-punctate and with dense silvery pubescence. Forewings smoky, the venation
black. Abdominal petiole flat dorsally and margined by gentle carinae, its sides
with a few irregular striae or carinae and with carinate ventral margins.

dé.—Length, 550 mm. Agreeing in colour and general characters with the
female. Funicle 1 a little longer than the scape; apical joint three-fourths as
long as funicle 1, one-half longer than the penultimate, pointed at apex. Abdominal
petiole slender, four times as long as wide; its dorsal surface with two parallel
carinae or striae which diverge at base and apex, the lateral surface with a few
strong irregular longitudinal striae or carinae.

Habitat—vVictoria: Beech Forest, three females, three males in January;
Millgrove, one male in November; all collected by F. E. Wilson.

Holotype and allotype in the collection of Mr. F. E. Wilson; paratypes in
the collections of the Queensland Museum, F.. E. Wilson, and the author.

M

THE SCIENTIFIC NAME OF THE COMMERCIAL OYSTER OF
NEW SOUTH WALES.

By Tom IREDALE and T. C. ROUGHLEY.

[Read 30th August, 1933.]

The common oyster of New South Wales has borne several scientific names
which will be discussed in a complete account of the oysters of Eastern Australia,
prepared by us but not yet published. This preliminary note is written to establish
the new specific name commercialis proposed for this species. It has been called
eucullata Born, introduced for a shell from Ascension Island and the West Indies;
it has been regarded as mordaz Gould, a ‘‘Feejeean” form; the name glomerata
has sometimes been pressed into use though that is strictly a Neozelanic species;
again circumsuta Gould has even been suggested, also described from the Feejees
and Samoa; while at other times subtrigona Sowerby,-.and also mytiloides Lamarck
have been advanced as substitutes.

OsTREA COMMERCIALIS, New name.

The name Ostrea commerciadlis is here proposed and a description given as
follows:

Normally this oyster is small, adherent, the lower valve rather deep, the
upper valve nearly flat, both valves radially crumpled. Coloration bluish
externally, whitish internally, juvenile specimens commonly showing radial
flames on a bluish ground.

The hinge-plate is of medium extent, the hinge-line short; edges of valves
internally more or less crenulated. The juvenile is flattened and nearly circular,
the ultimate shape depending on environmental stresses; with free growth, the
lower valve, which is attached, grows upward more or less regularly, forming a
deep cup with regularly crinkled edges. If growing in crowded situations the
growth becomes very irregular and often stunted, while on the open sea coast
it is commonly stunted and distorted owing to the prevailing high salinity which
retards growth-development. Under such conditions the internal edges tend to
develop rather strong teeth.

THE LIFE HISTORY OF THE AUSTRALIAN OYSTER (OSTREA
COMMERCIALIS*). .

By T. C. Roveutey, B.Sc., F.R.Z.S., Economic Zoologist, Technological
Museum, Sydney.

(Plates x-xxvii; two Text-figures.)

[Read 30th August, 1933.]

Introduction.

The history of the oyster in every country of its occurrence shows that at
some time or other the supply has fallen short of the requirements demanded by
commerce, and always scientific aid has been sought in an endeavour to account
for the depletion, and if possible to provide means for ensuring that a similar
shortage might be avoided in the future. Naturally, the propagation of the
oyster, involving the study of its life history from the fertilization of the egg
till the larval oyster attaches as a spat, when it comes under the control of the
cultivator, has on this account received extensive study. Embryological oyster
research had its origin in France, where very serious shortages occurred about
the middle of last century, at a time when science itself had made sufficient
progress to enable the problems attendant on such a subject to be investigated
with some degree of accuracy. Ever since that time the life history of the
oyster has been the object of specialized study by a large number of zoologists,
and a considerable amount of literature on the subject has accumulated. Mention
will be made here of a few only of the more important discoveries.

In 1852 the most notable advance up to that time was made by Davaine,
who found that the oyster of northern Europe (Ostrea edulis) was hermaphrodite,
being alternately male and female. In 1858 Coste made the first attempt to
develop oysters by artificial means, but his efforts proved to be entirely negative,
for the embryonic and early larval stages are passed in the mantle cavity of the
parent, and the artificial fertilization of the egg is rarely, if ever, successful.
It was demonstrated, however, by Bouchon-Brandely (1882) that the small
Portuguese oyster, O. (Gryphaea) angulata, was dioecious, and that fertilization
readily occurred when the eggs from the female and the sperms from the male
were artificially extracted from the parents and intermingled in sea water.

In addition to these workers, there have been many others who have assisted
in the elucidation of the problems connected with the embryology of the Huropean
species. Amongst these may be mentioned Lacaze-Duthiers (1854), Van Beneden
(1855), Mobius (1871, 1877), Horst (1882, 1884), Huxley (1883) and Hoek (1883).

The period 1883 to 1921 was one of little activity, but in the latter year
Orton began a series of researches which have elucidated many of the problems
unsolved by earlier workers. Notable, too, is the work of Sparck (1925).

It was not till 1879 that American scientists began to display an interest in
the life history of the American species (0. virginica). In that year Brooks
began a detailed study of the embryology of the oyster. It was he who first showed
that this species was dioecious, the eggs and sperms being ejected direct into
the water, where fertilization takes place, and the free-swimming stages of
development are undergone before the larva attaches itself and becomes a spat.

* This is the common commercial oyster of Australia for which the specific name
O. cucullata has usually been adopted (see page 278).

280 LIFE HISTORY OF THE AUSTRALIAN OYSTER,

Brooks has been followed by a long line of distinguished investigators, of
whom mention may be made of Ryder (1881), Rice (1883), Winslow (1884),
Jackson (1888), J. Nelson (1888-1917), and more recently by T. C. Nelson (1916-—
1932), Churchill (1919), Wells (1920), Prytherch (1928), Galtsoff (1930); and
in Canada, by Stafford (1905-1917).

Stafford’s researches are notable for their comprehensiveness, and his discovery
(1907) that the free-swimming period extends from three to four weeks in
Canadian waters was not suspected by most earlier workers. But previous
investigators had invariably tried to develop the young oysters through the
free-swimming period from artificially fertilized eggs, and endeavoured to deduce
the complete length of larval life from the time taken to reach the stage when
their cultivated larvae died. Stafford, however, by means of a plankton net towed
over the beds daily, was able to follow the development of the larvae under natural
conditions and watch the daily growth-rate till they settled down as spat. Having
obtained the larvae in every stage of growth, Stafford was afforded an opportunity
of studying their structure and movements at ages and sizes previously
unrecognized. Amongst other discoveries he found (1905) that in its later larval
life the oyster develops a foot by means of which it can crawl about quite
vigorously; he also identified a byssus gland at the base of the foot, and reasoning
by analogy with the function of such glands in other Lamellibranchs, he came to
the conclusion that in the oyster it produced the material which cemented the
shell to an object when the larva attached itself to become a spat. This was
shown by T. C. Nelson (1924) to be a correct deduction; he actually saw larvae
settle down on glass with the left shell valve held in contact with the substratum
by the foot while the secretion from the byssus gland was distributed by means
of the protruded mantle.

J. Nelson, in New Jersey, studied the daily growth-rate of normally occurring
larvae by filtering the water from the oyster beds. He found (1908) that the free-
swimming stage varied from two to three weeks according to the temperature.
Later (1915) he determined that at ordinary summer temperatures an interval
of from 13 to 16 days elapsed between the fertilization of the egg and the
subsequent attachment as spat. This has been confirmed many times by more
recent workers.

It has been the aim of practically every American investigator to raise spat
from artificially fertilized eggs, and several had announced that this had been
accomplished. The size to which they had developed their larve showed, as
Stafford points out, that in no instance had development proceeded far enough
to enable fixation to take place. The great trouble experienced by all workers
was filtration. A means had yet to be devised whereby a continuous flow ot water
would be permitted without losing the microscopic larvae. It remained for Wells
(1920) to accomplish this feat. He was able, by concentrating the larvae at
intervals in a small amount of water, by means of a milk separator, and then
transferring them to fresh sea water, to carry large numbers of larvae through to
the setting stage. In 1920 a thousand spat were obtained; in 1923, by improve-
ments in the technique, ten thousand spat were produced; in 1925 a total of a
hundred thousand spat was reached; and during 1926 a million larvae success-
fully set as spat.

Prytherch also, in 1923, succeeded in raising spat by artificial propagation.
He found that the problem of filtration was solved to a great extent by the use of
anew material known as “filtros”’, a white, rigid, porous, artificial stone, composed

BY T. C. ROUGHLEY. 281

essentially of silica, and similar to a sand filter in block form. The various
grades of porosity in which this material is made rendered it possible to retain
organisms of any definite size and still allow a good flow of water. For the
retention of the embryos a combination filter of loose, fine sand and filtros was
used until the embryo oysters had reached the shell stage; from that time on the
larvae were held in by the filtros blocks of various grades, which were changed to
suit the growth and size of the forms, and when the larvae were ten days old,
or older, they were held in by means of fine monel metal screens, which permitted
an unusually good flow of water. By this means large numbers of spat were
developed in from fifteen to twenty days from oysters which spawned naturally
on trays suspended just beneath the surface of the water.

The increasing success achieved by these experiments leads to the expectation
that eventually the artificial propagation of oysters may be carried out on a large
scale commercially, thereby eliminating the costly uncertainty which up to the
present has always attended the spatting season. Moreover, it is hoped that, by
selective breeding, the quality, shape and size of the oysters may be considerably
improved.

Of recent years considerable attention has been given to the oyster by
Japanese workers and much light has been thrown on the embryology of several
species occurring in Japanese waters, particularly on that of the common com-
mercial oyster, O. gigas. Prominent amongst these workers are Hori and Kusakabe
(1926), Amemiya (1928), and Seno (1929).

The Indian oyster (O. cucullata) received practically no attention until 1931,
when Awati and Rai published much valuable information on its structure and
breeding habits.

The world’s oysters may be divided into two main types: 1, those which
retain the embryos and larvae for a considerable time in the mantle cavity of the
parent, later ejecting the shelled larvae into the water where their larval develop-
ment is completed (larviparous type); and 2, those which spawn direct into
the water, where the fertilization of the egg is accomplished and where the whole
of the larval life is spent till attachment takes place (oviparous type). Examples
of larviparous oysters are O. edulis of Europe; O. lurida of the Pacific coast of
America; O. denselamellosa of Japan; O. angasi of southern Australia and
Tasmania; and O. lutaria of the South Island of New Zealand. Examples of
Oviparous oysters are O. angulata of Portugal (which has largely replaced the
indigenous O. edulis on the beds of France); O. virginica of the Atlantie coast
of America; O. gigas of Japan; O. cucullata of India; and O. commercialis of
Australia.

As long ago as 1852 it was shown that the larviparous European oyster
(O. edulis) was hermaphrodite, inasmuch as its sex was not constant, but
definitely changed at least from female to male. It has been shown by more
recent workers (Orton, Sparck) that during the course of its life, O. edulis under-
goes a series of sex changes, beginning as a male and alternating as male and
female throughout life, and described by Sparek as an alternating protandic
hermaphrodite.

In the case of the oviparous type, Brooks (1879) found that the oviparous
O. virginica was unisexual, each oyster remaining a male or female throughout
life. And since that date all oviparous oysters have been regarded as unisexual.
Kellogg, however, in 1890, recorded an individual of O. virginica whose gonad
contained both ova and sperms, and Amemiya in 1925 described three hermaphro-

282 LIFE HISTORY OF THE AUSTRALIAN OYSTER,

dite individuals of the oviparous Portuguese oyster (0. angulata). In 1928, the
writer recorded nine hermaphrodite individuals in O. commercialis (cucullata),
and was able to show for the first time that a sex-change occurs in this oviparous
species. Later in the same year Amemiya recorded cases of abnormal hermaphro-
ditism in the oviparous Japanese oyster (O. gigas), and the following year (1929)
showed that a sex-change occurs in this species also. Early in 1931 Awuati and
Rai recorded a sex-change amongst numbers of the Indian oviparous species
(O. cucullata).

Thus, during three years, three species of oviparous oysters, long regarded
as unisexual, have been shown to have a sex-change. But what of the other
oviparous species? The recorded occurrence of a hermaphrodite individual of
O. virginica, and of three individuals of O. angulata, raises the question whether
these species may not also change their sex. Further research may yet disclose
that no species of the genus Ostrea is unisexual.

It has been customary to regard the larviparous oysters as monoecious, and
the oviparous as dioecious, but in the light of recent research these terms are no
longer tenable, and it will save confusion if the terms ‘‘monoecious” and ‘‘dioecious’”’
are avoided.

In Australia the study of the life history of the oyster has received scant
attention. Tenison-Woods (1883) stated that the sexes are divided and the eggs
are probably discharged into the water where they may easily meet with male
cells. By mixing “the male and female fluid” it was found that fertilization of
the ova readily occurred.

Later, in 1888, Tenison-Woods appears to have altered his views concerning
the discharge of the sexual products into the water, and stated that “young
oysters are reared in the gill-chambers of the mother, in the case of the Australian
oyster, O. mordaz’. It seems probable that Tenison-Woods was confused in his
species, for the taxonomy of the New South Wales species was at that time very
unstable. In the first instance he was apparently dealing with the oviparous
O. commercialis, and in the second with the larviparous O. angasi, both of which
were then plentiful in the neighbourhood of Sydney.

Tenison-Woods believed that when expelled from the parent, the young were
sufficiently developed to be able to affix themselves as spat. He also stated that
the veliger stage possessed a ring of cilia and in the centre a long flagellum. He
was obviously unable to distinguish oyster larvae from those of other bivalves,
for at no period of its development does the oyster larva possess a flagellum,
though one is present in mussel larvae and forms a provisional byssus.

Altogether, Tenison-Woods’ observations were quite confusing and threw
little real light on the subject.

The work of Saville-Kent (1890), although it erred in a number of particulars,
contained much of value. He stated that in the case of O. glomerata (commer-
cialis) fertilization takes place in the water, “the young embryos . . . being
thrown upon their own resources from the earliest period of their existence’,
while the embryos of O. angasi are retained within the mantle cavity of the
parent, as are also those of the oyster of the south of New Zealand, which he
regarded as a local variety of O. angasi. Saville-Kent described the embryonic
development of O. commercialis as far as the complete envelopment of the
embryo by the shells, but erroneously assumed and actually illustrated their
attachment at that stage as spat, stating that the length of larval life occupied,
under favourable conditions, four days. His figure 21 shows the “earliest observed

BY T. C. ROUGHLEY. 283

attached condition of the oyster embryos or “spat”, attained to within the fourth
or fifth days succeeding the primary fertilization of the ovum; magnified about
50 diameters’, These so-called “spat” are shown 4:5 mm. wide; they represent,
therefore, individuals 0:09 mm. wide, or between one-third and one-fourth the
size of fully developed larvae or the earliest attached spat. Actually, they are
young shelled larvae of the straight-hinge stage, quite incapable of attachment.

Such, then, was the state of our knowledge of the life history of the Australian
oyster till the writer published a popular description in the Australian Museum
Magazine in 1925.

The present researches have been carried on intermittently since 1924. It is
hoped that opportunity will allow of their continuance, for much remains to be
done, and a particularly interesting problem awaits solution in the working out
of details of the sex-change. It is clearly shown that a change from male to
female occurs in the majority of oysters, but if is not known whether all of the
oysters change their sex, or whether any of them change from female to male.
I hope to have the opportunity of studying that problem in the near future.

The oyster with which this paper deals is the Australian edible oyster of
commerce. It appears to be confined to the eastern Australian coast, its range
extending from the far North Queensland coast to as far south as Wingan Inlet
in Victoria. In Queensland, the principal source of supply is in Moreton Bay,
where large quantities are grown and matured for market. Prolific crops occur
in Port Curtis and Sandy Bay, but they rarely thrive there owing mainly to a
too high salinity. Considerable quantities are transferred to Moreton Bay, how-
ever, where growth becomes much more rapid.

This oyster thrives best in those estuaries and rivers flowing to the east coast
of New South Wales which are fed by an abundance of fresh water from their
sources. The optimum water density lies between 1:015 and 1-020.

From New South Wales, where its cultivation is an important industry,
considerable quantities are exported to Victoria, South Australia and Western
Australia, its splendid keeping qualities allowing long distances to be travelled
without deterioration.

In Victoria, it occurs only in one or two rivers, principally Wingan Inlet,
on the extreme east, but there is no industry engaged in its cultivation in that
State.

The only other Australian oyster which enters into commerce is the larviparous
O. angasi, which resembles very closely the European O. edulis and the Japanese
O. denselamellosa. Small quantities only are marketed from the waters of
Victoria, South Australia and Tasmania.

The External Appearance of the Gonad.

Externally the gonad, when well developed, appears aS an enormous cream-
coloured organ extending from the anterior extremity of the animal, in close
proximity to the hinge, back as far as the pericardium. If the mantle is cut
away below the adductor muscle the gonad will be seen to continue along the oral
process to about two-thirds of its length. It is on the lateral walls of the oral
process that the gonoducts reach the exterior (Pl. x, fig. 1); they open into
slit-like depressions, one on each side. The same depressions serve as outlets
for the kidneys (organs of Bojanus), and they may therefore be designated
urino-genital clefts.

284 LIFE HISTORY OF THE AUSTRALIAN OYSTER,

In O. commercialis the cleft of the left side is situated a little behind the
anterior wall. of the adductor muscle (Text-fig. 1, w.g.c.); it lies in the middle
line; or a little above the middle line, of the oral process and crosses it almost
transversely, the dorsal edge being a little posterior to the ventral. On the right
side it is situated a little farther forward, and lies immediately beneath the peri-
cardium and in close proximity to the adductor muscle; on this side, also, it
crosses the long axis of the oral process transversely.

The disposition of the urino-genital cleft in this oyster varies deomewnnt from
that of O. edulis, An examination of several individuals of this species, sent to
me by Professor J. H. Orton, of the Liverpool. University, shows that the urino-
genital cleft opens longitudinally on the oral process on both sides in close
apposition with the adductor muscle (Text-fig. 1, A, B, u.g.c.).

Ucn

1 2

Text-fig. 1.—The situations of the urino-genital clefts of O. edulis and
O. commercialis compared. A. O. edulis, left side; B. O. edulis, right side;
C. O. commercialis, left side; D. O. commercialis, right side. a.m., adductor
muscle; 0.p., oral process; p., pericardium; w.g.c., urino-genital cleft.

Text-fig. 2.—Spermatozoon of Ostrea commercialis. x 1,500.

The gonoduct, as it extends forward on each side, immediately begins to
give off branches which coalesce and extend in all directions just beneath the
surface. In most ripe gonads these can be plainly seen; in some (PI. x, fig. 2)
they are very prominent, but occasionally an individual will be found with a
well developed gonad in which the ducts beneath the surface are practically
indistinguishable (Pl. x, fig. 3). Such a uniform gonad is frequently indicative
of maleness, and one with prominent ducts will usually be found to be an ovary,
but the sex cannot be judged with certainty by this means, for I have occasionally
seen males with extremely prominent ducts.

The gonads of the oyster are usually described as consisting of two in number,
one on each side, but so extensively do the ducts from one side coalesce with

BY T. C. ROUGHLEY. 285

those from the other side along the dorsal and anterior surfaces, and also the
follicles internally, that it is impossible to regard them as two entirely separate
organs. The disposition of the ducts shows that the bulk of the sexual products
produced on the left side find their way to the urino-genital cleft on that side, and
similarly with the right side, but it would be quite impossible to say through
which opening the genital products which are matured along the dorsal and
anterior regions of the gonad would emerge.

When the oyster begins to spawn the anterior region of the gonad is usually
the first to be drained (Pl. xi, fig. 4). After a partial spawning (Pl. xi, fig. 5)
the gonad appears as a very irregular mass; areas on each side may be completely
depleted, and others may retain the sexual products intact. If the spawning is
complete, the organs which were formerly obscured by the gonad, such as the
digestive diverticula and the intestine, become clearly distinguished through the
thin layer of vesicular connective tissue which covers them.

The Relationship of the Gonad with other Internal Organs.

The gonad is frequently spoken of as completely surrounding the digestive
organs; careful dissection, however, shows that this is not strictly correct, for
in the region where the digestive diverticula abut on the labial palps and gills
the gonad terminates on each side at the point of attachment of the mantles.
The relationship of the gonad to the organs which it partially encloses can be
studied quite effectively by slicing off with a razor sections about an eighth of
an inch thick, beginning at the anterior extremity and working backwards to
the termination of the oral process (Pl. xii, fig. 6). Examined with the aid of
a binocular microscope with a magnification of about 20 diameters, the disposition
of the principal organs can be clearly distinguished.

In the region nearest the hinge the gonads of the right and left sides are
confluent and appear as a homogeneous mass enclosed by a layer of vesicular
connective tissue which shows as a whitish band beneath the external epithelium.

In the region of the mouth, the digestive diverticula make their appearance
above the mouth, but are separated from it by a horizontal layer of the gonad.
Toward the anterior end of the oesophagus the digestive diverticula completely
surround it, and are themselves surrounded by the gonad above and on the sides;
the gonad, however, does not extend ventrally into the area between the oesophagus
and the labial palps. Towards the posterior extremity of the oesophagus, the
digestive diverticula no longer completely surround it; they are absent from the
dorso-lateral region on the left side. Here the gonad is separated from the
oesophagus by vesicular connective tissue only. Ventrally, the gonad on each side
terminates at the origin of the mantles, a large area of digestive diverticula
separating the gonads of the right and left sides, and lying in intimate contact
with the dorsal margins of the palps.

In a vertical plane above the anterior extremities of the gills, the digestive
diverticula are confined to the right side and to the horizontal area above the
gills. The gonad on the left side extends inwards almost to the oesophagus,
from which it is separated by a layer of vesicular connective tissue; it forms a
solid mass dorsally, and lies external to the digestive diverticula on the right side.
On the left side, and to a less extent on the right side, the gonad now begins to
invade the horizontal area above the gills.

In the region of the stomach the digestive diverticula occur in a dense mass
on the right side and below the stomach; on the left side they occur in islands

286 LIFE HISTORY OF THE AUSTRALIAN OYSTER,

bounded by vesicular connective tissue. Here the gonad still further penetrates
the area above the gills on both right and left sides.

Towards the posterior region of the stomach, the digestive diverticula do not
lie in such close apposition with the stomach wall, but are separated from it by a
layer of vesicular connective tissue which shows as a prominent band; they
enclose the stomach everywhere except dorsally. The gonad now almost completely
surrounds the digestive diverticula; the inwardly extending areas above the gills,
however, have not yet joined above the two inner hemibranchs.

Soon after the mid-gut leaves the stomach the digestive diverticula no longer
appear on the right side of the gut; they and the mid-gut are completely
surrounded by the gonad, the area above the gills now consisting of a continuous
mass of gonadial tissue. Within a short distance the digestive diverticula terminate
on the left side, and the gonad then forms a compact mass surrounding the mid-
gut and the returning loop of the intestine, and separated from them by a rela-
tively narrow area of vesicular connective tissue.

In the oral process, the mid-gut and returning intestine lie in the same
horizontal plane and in close apposition on the right side. In QO. edulis the
mid-gut lies above the returning loop of the intestine. The gonad on the left
side continues farther along the oral process than on the right side, and a little
distance beyond the urino-genital cleft.

Such are the principal relationships of the gonad with the digestive organs.
It is seen that it completely surrounds the digestive diverticula only in the
region behind the stomach. It is traversed by numerous blood vessels, the dorsal
vessel or anterior aorta showing prominently in all the sections cut anterior
to the pericardium except those closest to the hinge, where it divides into three
parts. Posteriorly, the anterior aorta lies in the centre of the mass of gonadial
tissue situated above the mid-gut, and as it travels forward it works slightly
to the left side, above the stomach and oesophagus. The posterior aorta becomes
conspicuous in sections in front of the pericardium where it is seen to descend
in close proximity to the mid-gut.

The loop of the intestine anterior to the stomach lies usually immediately
above the digestive diverticula, though occasionally it may traverse their upper-
most offshoots; it lies in the median vertical plane or slightly on the right side
until it begins to descend, when it crosses over to the left side and runs back-
wards in close apposition with the ventro-lateral border of the digestive diverticula.
It is embedded throughout its length in a layer of vesicular connective tissue.
This disposition of the intestine differs from that of O. edulis; in that species
the intestine, after leaving the oral process, crosses over to the left side and
lies above and to the left of the stomach, the anterior loop descending vertically.

The gonad when fully developed consists of anastomosing follicles with a
series of ducts on the surface, which serve to convey the ripe eggs or sperms
to the urino-genital cleft. In the very young oyster, in which the sexual products
are beginning to develop for the first time, the gonad consists solely of a series
of branching ducts which run in an antero-posterior direction; they lie very
close to the surface. The study of the disposition of these ducts can only be
done by a series of microscopic sections, and the description which follows has
been drawn from such a series in an oyster about six months old, which was
just beginning to produce sex cells. The sections were cut transversely, begin-
ning at the hinge end and terminating at the posterior edge of the mantle.

BY T. C. ROUGHLEY. 287

In sections of the anterior extremity of the body, the ducts form a complete
circle and are separated from the surface epithelium by a comparatively thick
layer of vesicular connective tissue, which, much condensed in proximity to
the external walls of the ducts, forms a continuous and fairly thick band external
to them. The area enclosed by the ducts is composed of vesicular connective
tissue only. In the region anterior to the mouth, above the dome-shaped anterior
extremities of the palps, the digestive diverticula make their appearance. The
ducts of the gonad surround them above and on the sides, but they do not
extend into the vesicular tissue which separates the digestive diverticula from
the mantle cavity.

In the plane where the mouth gives place to the oesophagus to form a hollow
tube compressed dorso-ventrally (Pl. xii, fig. 7), the digestive diverticula lie in a
compact mass in the region above the oesophagus, which is separated from them
by a thick layer of vesicular connective tissue. The ducts of the gonad do not
penetrate this layer, but terminate on each side above the attachment of the right
and left mantles, which in this region are confluent and form a sort of hood over
the palps.

As the oesophagus proceeds backwards and upwards it shortly enters the
region of digestive diverticula, which then come to surround it on all sides. As
the ducts of the gonad extend backwards, they gradually approach closer to the
external epithelium, until, about half-way along the oesophagus, they are separated
from it by a very thin layer of vesicular connective tissue which in places scarcely
exists at all; the ducts are now lenticular or elliptical in cross section, with the
long axis in some cases considerably produced. Their size varies greatly.

In the plane above the posterior extremities of the palps, or the anterior
extremities of the gills, which traverses the junction of the oesophagus and
stomach, the relative positions of the various organs have undergone little
alteration. The ducts of the gonad form a ring under the surface epithelium
above and on the sides, and still terminate above the junction of the mantles or
outer hemibranchs of the gills. A layer of vesicular tissue of irregular thickness
separates them from the digestive diverticula, except on the left side, where in this
region the digestive diverticula are absent. The ducts of the right and left sides
are separated inferiorly by a thick layer of digestive diverticula which occupies
the whole of the space between the stomach and the attachment of the gills.

In a vertical plane which traverses the anterior portion of the stomach, the
left kidney makes its appearance as a follicle situated above the junction of the
mantle with the body. It is shaped like an elongated duct of the gonad and lies
external to the lowermost of those ducts, between it and the external epithelium.
Proceeding backwards, other follicles of the left kidney soon appear and extend
inwards along the horizontal layer of vesicular tissue which occupies the region
between the digestive diverticula and the gills. In the middle region of the
stomach, the kidney follicles lie above the left mantle and the outer hemibranch
of the left gill. As the follicles of the kidney extend inwards they are accompanied
by the ducts of the gonad, which lie in the vesicular connective tissue between the
follicles of the kidney and the digestive diverticula. The ducts of the gonad now
vary very greatly in cross section; some are rounded in outline, while others are
oval or very elongated.

Toward the posterior extremity of the stomach, the ducts of the gonad begin
to invade the vesicular tissue above the mantle and gills on the right side; they
extend as far as the right outer hemibranch. On the left side they now extend as

288 LIFE HISTORY OF THE AUSTRALIAN OYSTER,

far as the left inner hemibranch; they are closely followed by the follicles of the
left kidney.

The follicles of the right kidney appear above the right outer and inner
hemibranchs of the gills in a vertical plane which traverses the anterior extremity
of the mid-gut and style sac; a follicle also appears now on the right side slightly
inferior to the right mantle on the dorso-lateral surface and external to the ducts
of the gonad; it is quickly followed by branches which extend about half-way
between the attachments of the right and left mantles dorsally and also on the
right side between the surface epithelium and the ducts of the gonad. Back
slightly farther they form an almost complete ring of follicles on the right side
extending from the dorsal aorta, midway between the attachment of the right and
left mantles dorsally, to above the attachment of the right outer hemibranch.
Dorsally, also, the ducts of the gonad curve inwards on both sides to lie below
the dorsal aorta and the hind-gut as it approaches the pericardium. Here, the
digestive diverticula occur only on the right and left sides of the mid-gut and style
sac; they no longer surround them above and below. On the left side the follicles
of the kidney now form a double layer. The ducts of the gonad continue to

penetrate the vesicular connective tissue above the attachment of the gills, and in
this region almost meet in the middle line.

In the region where the sections cut through the front of the pericardium
towards its dorsal aspect, a very different scene is presented. Dorsally we find
the rectum bounded laterally by the dorsal extensions of the mantle; below the
rectum lies the pericardium containing the ventricle. The follicles of the kidney
fill up in several layers the greatly thickened right wall of the pericardium and
extend below it in a single layer immediately below the surface epithelium to
above the right inner hemibranch of the gills. On the left side the wall of the
pericardium is extremely thin and no kidney follicles extend into it; they begin
dorsally immediately beneath the attachment of the left wall of the pericardium
to the oral process and extend down the lateral border of the oral process to above
the left inner hemibranch of the gills; they are particularly numerous in the
triangular area bounded on the left side by the surface epithelium, below by the
mantle and the outer and inner hemibranchs, and in the right dorso-lateral
region by the ducts of the gonad; they do not extend here into the dorsal region
of the oral process, below the pericardium. The ducts of the gonad form an almost
continuous layer along the dorsal surface of the oral process, below the
epithelium lining the pericardium ventrally, and on each side of the oral process
just internal to the kidney follicles. The mid-gut and style sac take up a consider-
able space in the centre of the sections; they are surrounded on the right and
left sides by digestive diverticula, which lie scattered in the vesicular connective
tissue and partially surround the loop of the intestine returning from the oral
process.

This disposition of the various organs continues till about midway beneath
the pericardium when the digestive diverticula no longer appear, the mid-gut and
accompanying style sac then being separated from the ducts of the gonad by a
thick layer of vesicular tissue only, except on the right side, which is traversed
by the returning loop of the intestine. Here also the right wall of the pericardium
contains but one extremely large kidney follicle; while on the left side a similar
large follicle has developed in the triangular area described above. These follicles
comprise what may be regarded as urinary chambers (right and left).

BY T. C. ROUGHLEY. 289

Below the posterior region of the pericardium and the auricles, the follicles of
the kidney again extend from each side into the dorsal surface of the oral process,
immediately beneath the pericardium, and are separated from the pericardium by
a thin membrane; they no longer extend into the ventral surface of the oral process
above the gills; they occur in greatest abundance and are largest in the right
wall of the pericardium, while on the left side they comprise the bulk of the tissues
between the style sac and the surface of the body bounded above by the pericardium
and below by the mantle and the left outer hemibranch of the gills. The ducts
of the gonad lie very close to the internal margins of the kidney follicles, and
these, too, no longer extend into the area above the attachment of the gills.

In sections cut through the posterior extremity of the pericardium, which
is here reduced to a relatively small cavity below the adductor muscle, the kidney
on the right side increases very greatly in extent; the urinary chamber now lies
in two planes like a letter L, the vertical arm extending from the adductor muscle
dorsally to immediately above the branchial cavity ventrally; while the hori-
zontal arm lies in the pericardium above the dorsal surface of the oral process,
from which it is separated by a layer of large kidney follicles. The left kidney
is here much smaller; the urinary chamber, irregularly oval in shape, with the long
axis vertical, lies in close apposition with the left wall of the oral process, while
follicles extend from its dorsal surface along the upper wall of the oral process
to meet those of the right kidney. Externally it is bounded by kidney follicles
which separate it from the surface epithelium above the attachment of the left
mantle, The ducts of the gonad are now small and few in number; they lie beneath
the dorsal surface of the oral process and extend on each side to the branchial
cavity.

It is in this region that the gonoduct of the right side opens into the urino-
genital cleft (Pl. xiii, fig. 8, u.g.c.). The duct lying nearest to the lateral wall of
the oral process is much larger than the rest; as it approaches the urino-genital
cleft it is crescent-shaped, the convex wall external and its surface strongly
ciliated, particularly in the region nearest the branchial cavity, while the internal
wall is lined by germinal epithelium and lies in close apposition with a prominent
nerve, the right visceral nerve (7.v.n.). The gonoducts of both sides are surrounded
by bands of sphincter muscle fibres immediately internal to the cleft, and are
lined by ciliated epithelium. The gonoduct of the right side runs in a dorso-ventral
direction; it is situated at the side of and slightly above the returning loop of the
intestine, which lies very close to the right wall of the oral process, the mid-gut
and style sac occupying the central area.

Slightly farther back the urinary chamber empties, by means of a short
canal, the ureter, into the urino-genital cleft. The ureter also runs in a dorso-
ventral direction, and is simply a ventral extension of the urinary chamber. The
ureter therefore opens slightly posterior to the gonoduct, and slightly to its right
in a somewhat higher plane; it is lined by ciliated epithelium, the cilia being long
and very numerous and filling up the lumen of the canal, The lower portion of
the urinary chamber is also lined by strongly ciliated epithelium, and is surrounded
by a narrow band of muscles which extends to the ureter and encloses it on all
sides. The right visceral nerve, which lay in close contact with the genital duct,
here lies internal to and below the wall of the ureter, while immediately external
to the wall of the urinary chamber, a little above the ureter, lies the larger right
branchial nerve (0D.n.).

290 LIFE HISTORY OF THE AUSTRALIAN OYSTER,

The gonoduct on the left side opens into the urino-genital cleft (Pl. xiii,
fig. 9, u.g.c.) slightly posterior to that of the right side; it crosses the median line
of the oral process and, like that of the right side, extends in a dorso-ventral
direction. Its internal surface is in close contact with the left visceral nerve
(lw.n.).

A little farther back again the ureter opens downwards into the urino-genital
cleft. The urinary chamber on the left side lies alongside the oral process, and
not dorsal to it as on the right side. The urino-genital cleft is bounded internally
by the epithelium of the oral process which is here ciliated, and externally by
similar ciliated epithelium which is separated from that of the urinary
chamber by a thin layer of connective tissue. As on the right side, the walls of
the urinary chamber and ureter are surrounded by a thin band of muscle fibres.
Immediately above the ureter the left visceral nerve runs in an antero-posterior
direction. Ventrally the wall of the urinary chamber is traversed by the large left
branchial nerve (b.n.) which causes a pronounced dilatation of the wall. The
epithelium of the urinary chamber is lined by cilia which become very long and
numerous in the vicinity of the ureter.

The ducts of the gonad lying just beneath the dorsal surface of the oral
process extend a little farther back than the openings of the gonoducts on each
side, and terminate at about the level of the ureter.

In the plane of the left ureter the follicles of the kidney form a thick layer
between the adductor muscle and the dorsal surface of the oral process along its
whole length; on the right side they do not extend below this level; on the left
side, however, they continue downwards on the side of the oral process as far as
the opening of the ureter.

Continuing posteriorly, the kidney follicles extend beyond the termination of
the oral process to about the posterior surfaces of the visceral ganglia.

The Development of the Gonad.

In the very young oyster, in which the sexual products are developing for the
first time, the layer of vesicular connective tissue surrounding the digestive
diverticula is of very irregular thickness; in places the digestive diverticula may
extend almost to the surface epithelium, while in others the intervening layer of
vesicular tissue may be of considerable thickness, but nowhere does it attain the
great depth characteristic of an older oyster which is about to develop sexual
products at the beginning of the summer. In this young stage all oysters from
which sections have been prepared were males, and it is, I think, very unlikely
that the development of the eggs can be studied in an oyster of this species which
has not previously developed spermatozoa. Just beneath the surface epithelium,
and separated from it by a layer of vesicular tissue from one to three or four cells
deep, the ducts of the gonad, mostly elongated in cross section and lenticular in
shape but occasionally smaller and broadly oval, extend in a band round the oyster.
These ducts are lined along their outer surfaces by ciliated epithelium, and along
their inner surfaces by germinal epithelium which, in the youngest oysters of
which I have prepared sections, is already giving rise to spermatogonia; these,
however, have not yet begun to produce spermatocytes. Irregularly distributed
amongst the small, flattened cells of the germinal epithelium are some, the
spermatogonia, which have become relatively greatly swollen and contain a large
nucleus with a great number of chromatin granules. These spermatogonia are

BY T. C. ROUGHLEY. 291

irregular in outline and variable in size, the largest being 144 in diameter and
their nuclei 10u.

In slightly older oysters proliferation of the spermatogonia into spermatocytes
and development of the latter into spermatozoa can be clearly distinguished
(Pl. xiv, figs. 10, 11). The primary spermatocytes, or spermatocytes of the first
order (Pl. xiv, fig. 10, sp.1), form a layer immediately above the spermatogonia
(spg.); they are rounded in shape and measure up to 6u in diameter; the nucleus
occupies the bulk of the cell, and its chromatin granules, although numerous and
deeply stained, are somewhat scattered, and give to the nucleus a general appear-
ance of being lightly stained. Immediately above the primary spermatocytes, the
secondary spermatocytes, or spermatocytes of the second order (sp.?) lie in a
somewhat deeper layer; these measure up to 3:54; the nucleus is large, and the
chromatin more concentrated, hence they appear to be more deeply stained. Above
the secondary spermatocytes, and almost completely filling the lumen of the duct,
are the spermatids and spermatozoa (s.). The heads of the latter measure 1:5y in
diameter (in sections), and their chromatin, now extremely concentrated, stains
very deeply. The tails lie in bands between the lanes of spermatozoa, and always
point towards the outer surface of the duct.

As development of the testis proceeds, the vesicular connective tissue, both
external and internal to the genital ducts, increases rapidly till it forms a thick
layer separating them from the surface epithelium externally and from the
digestive diverticula internally. When the ducts are distended with spermatozoa,
offshoots begin to sink into the vesicular tissue which borders them internally.
These may extend in irregular follicles as far as the digestive diverticula, or they
may give off branches which, running more or less horizontally, are cut across
in sections made at right angles to the surface of the gonad (PI. xv, fig. 12).
Germinal epithelium usually lines the walls of these follicles, from which
spermatogonia, spermatocytes and spermatozoa continue to develop. Nourishment
for the developing sex cells is obtained from the vesicular tissue, and as
proliferation proceeds the ducts and follicles tend to coalesce with the partial or
even total disappearance of the follicular walls, until, as the testis approaches
maturity, they may almost cease to exist as separate entities.

In a male oyster just prior to spawning (Pl. xv, fig. 13), the vesicular tissue
lying between the genital ducts and the surface epithelium may become reduced to
a narrow band, and in places may disappear entirely; the ducts in cross section
are of great width and are divided from one another by narrow bands of vesicular
tissue which extend downwards into the testis for varying distances, becoming
narrower as they proceed, while the spermatozoa lie in intimate contact with
the ciliated epithelium of the ducts. The testis rarely becomes a uniform mass of
spermatozoa, but is broken up into irregular islands by small areas of vesicular
tissue, which are frequently bordered by spermatocytes. In these areas of vesicular
tissue an occasional blood space is to be found. The tails of the spermatozoa are
usually bunched together in definite streams, which run in all directions.

The head of the live sperm of O. commercialis measures 2u in diameter, and
the tail 42u long. The shape of the head of this sperm differs from that of
O. virginica, which is described by Stafford (1913) as “almost oval in form, some-
what pointed anteriorly, but inclined to be squarish posteriorly’. The head of the
sperm of O. commercialis is spherical (Text-fig. 2). Attached to the head, and
grouped round the insertion of the tail are four spherules, of which usually only
two, occasionally three, can be seen when the sperm is lying on its side.

292 LIFE HISTORY OF THE AUSTRALIAN OYSTER,

It need scarcely be remarked that sperm morulae, characteristic of larviparous
oysters, do not occur in O. commercidlis.

The head of the sperm of the American oyster, O. virginica, is stated by
Stafford (1913) to measure 1-754 in breadth, and the tail 354 in length; while
Churchill (1920) states that the head measures 2-5u, and the tail about 50y; that
of O. edulis is stated by Hoek (1883) to be at most ly, and the tail 25u, while
Orton in the Encyclopaedia Britannica (14th Edition) gives the measurement of
the head as 2u, and the tail about 20-30y.

When considering the development of the ovary, it should be borne in mind
that we are dealing with an oyster which has probably already functioned as a
male. In all young females which I have examined by means of microscopic
sections, the layer of vesicular connective tissue in which the germinal ducts lie
is of far greater depth than in young males in which the sex organ is developing
for the first time. This vesicular tissue (Pl. xvi, fig. 14, v.t.) forms a thick layer
between the ducts and the surface epithelium, and also between them and the
digestive diverticula. There is, therefore, already awaiting the development of
the eggs, which require more nutriment for their growth than the sperms, an
abundance of vesicular tissue in which nutriment, mainly in the form of glycogen,
is stored.

External to the genital ducts the layer of vesicular tissue is usually homo-
geneous and of rather loose texture; occasionally, however, patches occur in which
the tissue becomes very condensed, the cells losing their characteristic form of
bladder cells and becoming smaller and more compact. Similar condensed areas
may occur in the broader band of vesicular tissue lying internal to the ducts; in
the latter, numerous blood spaces occur, the tissue bordering them being usually
condensed.

The germinal ducts (Pl. xvi, figs. 14, 15, d., d.1) are for the most part lenticular
in. cross section, but oval or circular ducts are not uncommon. The distance
between the ducts varies greatly; in some cases a band of condensed vesicular
tissue of variable width extends from one duct to the next; in others, it may
continue along the external surfaces of the ducts and form a more or less unbroken
band bordering them externally (Pl. xxii, fig. 26, v.t.*); in other cases, again, no
condensation at all may occur in the tissue between or external to the ducts.
Muscle fibres are frequently present in this band of tissue, particularly in that
portion of it in contact with the ciliated epithelium of the ducts.

As in the testis, the ducts are lined by ciliated epithelium externally (Pl. xvi,
fig. 15, Pl. xvii, fig. 16, c.e.), and internally by germinal epithelium, from which
the ova develop. In their very earliest stages it is extremely difficult, if not
impossible, to distinguish the male from the female germ cells. There is usually
no difficulty, however, in determining the sex of the individual, for easily recogniz-
able young ova early become apparent in some of the ducts; they are characterized
by their granular cytoplasm, large, comparatively clear nucleus, and conspicuous,
deeply-staining nucleolus. As they develop they become enormously swollen,
attached to the basement membrane by a broad base, and project into the lumen
of the duct as a dome-shaped cell, the nucleus lying close to the attached surface.
As they continue to grow, the development of other ova at their bases compresses
the region of their attachment till it becomes constricted into a long, narrow neck,
the free portion remaining swollen, and the whole resembling an attenuated pear
(Pl. xvii, fig. 16, o.). The nucleus now lies in the centre of the swollen extremity,
with the nucleolus close to its circumference.

BY T. C. ROUGHLEY. 293

With the crowding of the germinal layer by the rapid development of the
eggs, portion or portions of it evaginate into the vesicular tissue, and, by further
evaginations, form numerous branching and anastomosing follicles (Pl. xvi,
figs. 14, 15, f.). As the ova in these follicles develop, they are continually absorbing
nourishment from the vesicular tissue, which rapidly diminishes in extent, till in
the ripe ovary, none, or practically none, remains. The mature ovary, therefore,
consists of a mass of closely packed ova attached to the follicular walls by their
narrow extremities (Pl. xvii, figs. 16, 17, 0.). The germinal ducts, in the mean-
time, have become greatly extended laterally; many have joined up with their
neighbours, whilst others are separated only by a narrow band of vesicular tissue
or by the walls of the follicles. The layer of vesicular tissue lying between the
ducts and the surface of the oyster also diminishes rapidly as the eggs develop,
till in the mature ovary it becomes reduced to a very narrow layer (PI. xvii,
fig. 16).

Conspicuous amongst the cells of the ciliated epithelium of the germinal ducts
of both the ovary and testis are frequently to be seen granular cells which stain
very deeply with eosin and other acid stains. These cells may be relatively small,
no larger than the epithelial cells, but more frequently are greatly swollen and
project well into the lumen of the duct (PI. xiv, fig. 11; Pl. xvi, fig. 15; Pl. xxiii,
fig. 28, s.c.). Occasionally neighbouring cells may coalesce to form a continuous
layer of considerable extent. Some ducts may be entirely devoid of them, whilst
in others the ciliated epithelium may be entirely replaced by them. They may be
seen at times extruding their granular contents into the lumen of the duct.
Whilst these granular cells are usually devoid of cilia, occasionally a few cilia
may be seen projecting from the surface; they appear, therefore, to have originated
as ordinary epithelial cells, later differentiating to secrete granular material.
They occur in the ducts of the gonad in all stages of its development, from the
time when the sex cells are beginning to differentiate, till the gonad is mature.
They reach their greatest development, however, and usually occur in greatest
abundance, in partially spawned or completely spawned oysters (PI. xviii, fig. 19;
Pl. xix, fig. 21; Pl. xxi, fig. 24, s.c.); in several successive ducts they may
completely replace the ciliated epithelium, and form a layer from one to several
cells deep, and may even fill almost the whole of the lumen of the duct. But in an
oyster in which this excessive development occurs, a few of the ducts may remain
almost, or even entirely, devoid of them. Such prolific development, however, is
not to be found in all recently spawned oysters, for occasionally one will be found
in which the granular cells occur in very few of the ducts, and are inconspicuous
and sparsely distributed.

The function of these cells I have been unable to determine. They are not
mucous cells, mucicarmine having no affinity for them whatever. They stain
intensely with eosin, erythrosin, acid fuchsin and iron haematoxylin, and very
slightly with Delafield’s and Ehrlich’s haematoxylin. In preparations stained with
iron haematoxylin and acid fuchsin, the granules in some of the cells are rendered
a deep bluish-black, whilst in others they are a brilliant red. In eosin-methylene
blue preparations they form a beautiful contrast.

Granular cells with similar staining characteristics are found in the surface
epithelium, where at times they occur in great abundance and may even form
an almost continuous layer of considerable extent, and in that of the intestine.
In the latter situation they have been noted by several authors in various molluscs,

N

294 LIFE HISTORY OF THE AUSTRALIAN OYSTER,

and have usually been designated “eosinophilous cells’ from their great affinity
for this stain.

Mature eggs of the oyster may be studied either alive or in sections; in the
former case, they may be obtained by pricking the surface of a ripe ovary, or
preferably by a little gentle pressure of the finger along the surface of the
ovary from before backwards, thereby forcing some of the eggs out of the
oviduct. The eggs of O. commercialis (Pl. xviii, fig. 18) may be round, oval,
or more commonly pear-shaped; if the latter, the narrow end may be greatly
prolonged. They vary considerably in size; the width of pear-shaped eggs varies
usually from 0:05 to 0:06 mm., while the longest I have seen measured 0-225 mm.
The largest eggs I have measured were subspherical, the diameter being 0:09 mm.
The shape of the egg is not an indication of maturity, for an elongated pear-
shaped egg is as capable of fertilization and subsequent development as a spherical
one. Soon after fertilization, however, an elongated egg becomes spherical. In
the Bombay oyster (0. cucullata), Awati and Rai (1931) found that the spherical
eggs alone were ripe and capable of fertilization.

The eggs of all oviparous oysters appear to average approximately the same
size, i.e., 0°05 mm., as stated by Stafford (1913), Churchill (1920), and T. C.
Nelson (1921) for the American O. virginica; Amemiya (1928) for the Japanese
O. gigas, O. spinosa, and O. circumspicta; and Amemiya (1926) for the Portuguese
O. angulata. The eggs of the larviparous types are all larger and measure 0-1 mm.,
as stated by Amemiya (1928) for the Huropean O. edulis and the Japanese
O. denselamellosa; and Stafford (1918a) for the American O. lurida.

The eggs are enclosed in a very thin membrane; they contain granular
cytoplasm with a narrow, clear, homogeneous band immediately beneath the
lining membrane, and another surrounding the nucleus. The nucleus is very
large and spherical in shape; it contains granular nucleoplasm more transparent
than the cytoplasm surrounding it. The nucleolus appears as a small glistening
spherical object, which usually lies in close apposition with the nuclear membrane,
but may occur anywhere in its interior.

In sections stained with haematoxylin the granules of the cytoplasm, the
nuclear membrane, and particularly the nucleolus, stain deeply, while the proto-
plasm of the nucleus stains lightly and tends to become reticulated (PI. xvii,
fice Os)

The Gonad during and after Spawning.

When an oyster spawns, the reproductive elements, ova or sperms, are con-
ducted along the ciliated ducts to the gonoducts on each side, which, as we have
seen, open on the sides of the oral process beneath the adductor muscle. As
batches of eggs or sperms are swept along these ducts, further batches are drawn
upwards, where they enter the stream formed by the beating of the cilia. In this
way the follicles are gradually drained.

Although I have prepared sections of many oysters during and after the
spawning period (during late summer and winter), I have not found one in which
spawning has been complete, i.e., in which the whole of the eggs or sperms have
been ejected. It is apparently usual for some to remain in the gonad; indeed, at
times the bulk of them are retained, and individuals are occasionally found in
batches of recently spawned oysters which have obviously not spawned at all. In
the latter case the eggs or sperms may either be carried over the winter or, as
appears to be more frequent, they may be absorbed.

BY T. C. ROUGHLEY. 295

Absorption of the undiscarded eggs or sperms of a partially spawned oyster
appears always to take place; the process is a comparatively slow one, for various
stages of absorption may be seen during the whole of the winter following
spawning.

In the case of the testis, the ducts and follicles containing spermatozoa become
infiltrated with small, irregular, densely-crowded, vesicular cells (Pl. xviii, fig. 19,
v.t, v.t.2). These cells are in very active division and mitoses are greatly in
evidence. Gradually, as the spermatozoa are absorbed, they are completely
replaced by the condensed vesicular cells (PI. xix, fig. 20, v.t., v.t.2), which continue
to develop and expand until, early in the following spring, they assume the
typical bladder-like character of the ordinary vesicular cells characteristic of this
region. Their function is, of course, to store nutriment for the ensuing develop-
ment of spermatozoa or, possibly, eggs.

In the case of the ovary, absorption of undischarged eggs takes place in a
similar manner. A few small vesicular cells make their appearance amongst the
eges in the ducts and follicles, and develop at the expense of the eggs (Pl. xix,
fig. 21; Pl. xx, fig. 22, o., 04, v.t1, v.t2). Gradually, as the vesicular cells increase
in numbers, the eggs degenerate, decrease in size, and eventually disappear
altogether, their place being taken by densely crowded vesicular cells (Pl. xx,
fig. 23; Pl. xxi, fig. 24, v.t4, v.t2). When absorption is complete the vesicular cells
continue their growth by extracting nourishment from the blood stream, until the
whole of this region comes to consist of large, more or less uniform, vesicular or
bladder cells which characterized it before the sex cells began their development.

As the genital products are absorbed, the ducts shrink away from their
neighbours with which they had coalesced as the gonad reached an advanced
stage of development. They remain filled for a considerable time, however, with
the condensed vesicular cells (Pl. xxi, fig. 25, v.t2) which have replaced the eggs.
Gradually these cells migrate downwards until the ducts are completely free from
them; the ducts then again assume the form they had when the sex cells began
to differentiate, i.e., they are lined by germinal epithelium internally and ciliated
epithelium externally.

Considerable difficulty was encountered in the preparation of sections of
recently spawned oysters. The difficulty was apparently one of fixation, for the
layer of vesicular connective tissue in which the sex cells were embedded, and
sometimes also the layer between the genital ducts and the surface epithelium,
was usually found to undergo considerable disintegration during spawning. The
fixatives used consisted of Bouin’s picro-formol-acetic, Zenker-formol (Helly’s
fluid), and Carnoy’s acetic-alcohol-chloroform mixture. None of them was found to
be satisfactory. For general histological work on the oyster, I have found Bouin’s
fluid to be a satisfactory fixative; Zenker-formol was found to distort some of the
tissues badly; and Carnoy’s fluid, excellent for some tissues, was found very poor
for others; it usually caused a shrinkage of the ducts and the breaking away of
the cilia from the epithelium. In my earlier histological work most of the tissues
were fixed in formol-alcohol-acetic of Bles, with fairly satisfactory results.

The Spawning of O. commercialis.

My observations on the spawning habits of O. commercialis were made at Port
Macquarie, a harbour at the entrance of the Hastings River, on the north coast of
New South Wales. Port Macquarie is a large basin which receives the water from
the Hastings River at its north-east corner, and has a narrow outlet to the Pacific

296 LIFE HISTORY OF THE AUSTRALIAN OYSTER,

in the south-east quarter through which the tidal waters reach the sea. Owing to
the narrowness and shallowness of this channel, the rise and fall of the tide in the
estuary is never very great. The channel itself is bounded along its southern
border by a breakwater, and requires frequent dredging to allow of the passage
of craft of even shallow draught. The harbour has two channels, the eastern being
navigable by steamers of small draught, while the western is impassable by ocean-
going vessels of any description, the central area consisting of a large sand spit
which is bared by the tide.

Oysters are cultivated along the whole of the western foreshores to within
about half a mile of the breakwater. Choice of this locality was made because
the water is usually very clear, allowing of unobscured observation of the cysters
when submerged, and also because the actual spawning is usually heavy, and
abundant catches of spat are a general rule. Owing to the proximity to the ocean
the density of the water is usually rather high, though a good deal of fresh water
spills over from a large swamp inshore after heavy rain.

Having received, about the middle of February, 1924, some oysters from Port
Macquarie which had the reproductive organ in a very advanced state of develop-
ment, containing an abundance of ripe ova and sperms, I proceeded there on
25th February and made a daily survey of the beds in an endeavour to discover
any indications of spawning oysters; in this I was assisted by the local Inspector
of Fisheries and by the oyster growers, who were working the beds continuously.
Although the oysters everywhere were very fat, no spawning was seen. until
5th April, and although numbers were repeatedly opened for examination, none
previous to this date had given any indication of even a partial spawning. In
the meantime a plankton net consisting of No. 16 bolting silk (157 meshes to the
inch) was towed over the beds daily, in many situations, at different states of the
tide, and from the surface of the water to close to the bottom, but in no instance
was an oyster larva seen, although the larvae of many other bivalves were always
abundant (Pl. xxiv, fig. 30).

On 5th April, the late Mr. Thomas Dick, a prominent oyster grower of the
district, noticed oysters beginning to spawn a short distance from where my
laboratory was situated. Hastening to the bed, I found that the water over it was
distinctly cloudy, and in a very short time the bottom and the stones to which the
oysters were attached were obscured from view in water two feet deep. Spawning
had begun about 12.30 p.m.; the temperature of the water at the surface was 70° F.,
and at the bottom 68° F.; the density at the surface was 1-0195, and at the bottom
1:020; the tide, which had been a particularly high spring tide (new moon) was
from three to four hours on the ebb, and when the oysters began spawning was
about mean high water level. The wind was moderately strong from the south,
though the beds were fairly sheltered by a lee shore; the tide was ebbing very
slowly, a heavy sea outside the bar tending to bank the water in. Heavy rain had
fallen during the previous day, but the density of the water was little affected by
it. By 1.30 p.m. (an hour after the commencement of spawning), the water had
practically cleared, and although the beds were patrolled for about a quarter of a
mile, no evidence of spawning could be seen beyond the bed under observation,
which covered an area of about a hundred yards long by about twenty wide.

The late Mr. Thomas Dick, whose observations in the realm of oyster life were
particularly keen and very reliable, informed me that spawning invariably occurs
on these beds during spring tides when two or three hours on the ebb, and often

BY T. C. ROUGHLEY. 297

when a heavy sea is running outside. On one occasion previously he had taken
the temperature of the surface water over a shallow bed and found it to be 70° F.

The temperature of the water over the beds had been taken daily from
25th February till spawning occurred on 5th April, and on only one occasion was it
found below 70° F., when on the day preceding spawning it was 68° F. It varied
from 70° F. to 85° F., and was usually from 72° F. to 76° F. The oysters during the
whole of this time appeared to be capable of spawning, and fertilization was always
readily effected when ova and sperms were mixed in sea water. It would appear,
therefore, that other physical conditions besides temperature play a part in
governing the spawning impulse.

There is abundant evidence that the heaviest spawnings occur on the New
South Wales coast at spring tides (both full and new moon tidal periods), and
cultch is frequently laid out just before Christmas, when a heavy spawning often
occurs during the unusually high tides which occur at that period. But if spawning
were confined to spring tides the resulting sets of spat would assuredly occur
in well defined fortnightly pericds, for, even allowing for irregularities of growth,
the bulk of the larvae may be expected to develop with a certain amount of
uniformity. An investigation during three months on the Hawkesbury River in
1925, however, showed that this, at least in that river, is not always the case.
The attachment of spat occurred with the utmost irregularity; practically every
day a few spat were found attached to shells, fibro-cement sheets, glass, and other
objects placed out to secure them. On some days more would attach than on
others, but there was never a heavy set during any period of twenty-four hours.
In the aggregate, the catch was a good one, but it extended over a period of five
months. It appears to be a fair assumption that spawning in various localities was
proceeding daily or almost daily during the whole of that period. Such conditions
appear to be normal to the Hawkesbury River; inquiry amongst the most
experienced cultivators showed that none had ever seen the water rendered milky
by oyster spawn, and few indeed had ever seen a spawning oyster. Apparently
it is customary for the oysters to spawn partially at intervals, and in this respect
they are in marked contrast with those at Port Macquarie, where a general and
heavy spawning may occur during the course of one tide, resulting in an almost
simultaneous set of spat. I have seen fibro-cement sheets from Port Macquarie
with an enormously heavy set of very young spat which had every appearance
of being of a uniform age. Oyster growers there have seen the water clouded with
spawn for hundreds of yards.

Further observations of the spawning of the oysters on 5th April, 1924, showed
that two methods were commonly employed to expel the genital products: one,
the more common, being a forcible ejection, the spawn in this case leaving that
part of the shell situated closest to the gonoduct, i.e., directly inferior to the
adductor muscle; and the other, a mere dribble, when the spawn usually oozed
from the shells in the exhalant current at the dorsal surface, above and behind
the adductor muscle; less frequently it would ooze from the same region
(inferiorly) as that from which it was forcibly ejected.

The forcible ejection was accomplished by a gradual opening of the shells,
followed by a sudden closure, accompanied, when out of water, by a distinct hissing
sound, the spawn leaving the shells in a cloud, which resembled the form of the
smoke emitted when a cannon is fired. Immediately after leaving the shell it was
concentrated in a thin, swiftly-moving stream, but gradually became more diffuse

298 LIFE HISTORY OF THE AUSTRALIAN OYSTER,

and slowly dispersed into the water. The intervals between each ejectment were
of short and irregular duration during the early stages, but gradually became
longer as the process was continued; the force of the expulsion was also greater
at the beginning than towards the end.

It has been stated by T. C. Nelson (1921) that the spawn of the American
oyster oozes from the gaping shells of the males, while the females forcibly eject
it at intervals, and a similar observation was made by Prytherch (1923). At Port
Macquarie no such rigid differentiation could be made, for sperms were occasion-
ally seen to be forcibly ejected from the male oysters, and eggs sometimes to
trickle from the females, in each case from the ventral margin of the shell.
Observations were made on specimens which were placed singly in glass jars, and
verification in each case was obtained by means of microscopical examination.
It was also seen that the spawn may be forcibly ejected or ooze out of the same
oyster, though the exudation in this case was only seen after the oyster had been
spawning for some time. After one ejection the shells remain closed for some
time; they then slowly open very widely, as much as a quarter of an inch, when
the sudden snapping of the shells is repeated.

A stone with oysters attached was removed from about two feet of water to
shallower water inshore where portion only of the stone was submerged; here
the oysters both in the water and exposed to the air continued to spawn; in the
latter case one squirted the fluid into a beaker held at the level of the oyster
about twelve inches distant.

A marked contrast was noted between the dispersal of the eggs and that of
the sperms. The eggs quickly diffused, whereas the sperms were inclined to hold
together for a short time as well defined white streaks. Numbers of oysters, having
begun to spawn in the water, were lifted into the boat, and in almost every case
they continued to spawn at intervals similar to those remaining in the water.
Several, having completed spawning, remained with the shells gaping widely; these
were left in the boat and two days afterwards were found to be dead, although
normally this species of oyster will live for upwards of two weeks out of water.
The exhaustion of spawning appeared to render them utterly unable to effect the
closure of the shells, and in the absence of a normal flow of water, they quickly
perished. Several of the gaping oysters were opened and in most cases it was
found that about half the contents of the gonad had been drained. The extent to
which spawning continues appears to regulate the degree of exhaustion.

Having begun to spawn, the oyster will continue the process in spite of rough
handling. For instance, some oysters which had been seen to eject spawn were
forcibly knocked, by means of a culling iron, from the stone to which they were
securely attached, and when placed in a jar of water soon opened their shells and
continued to eject the spawn.

If spawning occurs on a bed at a time when they are being gathered for
market, it is necessary to return all the oysters to the bed, for they may continue
to spawn in the bag and die before they reach the market.

A marked difference was noticed in the nature of the fluid ejected from the
oysters at the beginning of spawning and when it was nearing completion; in
the former case it was more uniform and dispersed more rapidly, while in the
latter, it was accompanied by relatively large pieces of tissue, apparently from
the broken-down gonad.

The eggs and sperms quickly settle to the bottom. In a large beaker into which
four spawning oysters were placed, the water was for a time quite milky and

BY IT. C. ROUGHLEY. 299

Opaque, and the outline of the oysters could not be seen when looked at from
above; settlement was proceeding the whole time, the water at the bottom of the
beaker becoming increasingly opaque, while that at the surface became gradually
clearer, At the end of half an hour the water was perfectly clear, and the spawn
lay in a dense layer on the bottom of the glass and on the surface of the oysters
like a mantle of snow.

In an endeavour to procure as much information as possible relative to the
state of the tide when O. commercialis spawns, I called for reports (through the
courtesy of State Fisheries) from the Inspectors of Fisheries stationed along the
New South Wales coast. These Inspectors are continually moving about the oyster
leases, and have a splendid opportunity of observing the condition of the oysters.
Altogether, 22 instances of spawning were reported, and of these 15 were to the
effect that spawning occurred on the ebb tide, and 7 on the flood. Eleven spawnings
were recorded during spring tides, and four during neap tides; in seven of the
spawnings reported no note had been taken of the tidal range.

Three of these reports drew attention to the fact that young mullet (Mugil sp.)
were attracted to the spawning, and by their activity, they gave every indication of
devouring large quantities of spawn.

In two instances the Inspectors stated that they had seen the spawning
continue after the oysters were bared by the tide, leaving the shells more or less
covered with the white fluid.

For purposes of comparison it will be of interest to survey the observations of
the spawning of different species recorded by various authors in other parts of
the world.

Orton (1920) states that, throughout its range, the larviparous Huropean
oyster (0. edulis) appears to begin to breed at a temperature of about 59° F.—61° F.
in varying salinities from about 25%, to as high as 36%, or more in the Gulf of
Naples, and continues to breed so long as the temperature remains above this
figure, so that in the warmer situations there is a longer breeding period than in
the colder ones.

Sparck (1925) found, however, that the breeding temperature varies according
to whether the spring rise in temperature is rapid or slow. The breeding
temperature was found to be higher the more steeply the temperature curve rises.
After a warm spring, oysters were found with larvae in the mantle cavity at a
temperature of 55° F.-57° F.; after a cold spring, when the temperature of the
water rose rapidly, no oysters with developing larvae were found till a temperature
of 61° F.—63° F. was reached.

A lunar periodicity has been stated by Orton (1926) to occur in O. edulis, a
maximum of spawning being found round about the days of full moon. It is
suggested that the rapid rate of change in pressure accompanied by increased
temperature at spring tides may be sufficient stimulus to cause spawning to occur.
In the Fal estuary in 1926, Orton (1927a) found that regular spawning began on
the spring tides at about full moon at the end of June, and continued throughout
the summer until the full moon spring tides about 23rd September.

In the American oyster (0. virginica), J. Nelson (1890) found that in New
Jersey spawning begins when the temperature of the water reaches 70° F.

Stafford (1913) found that in Canadian waters the temperature of the water in
the region of oyster beds at the beginning of spawning approximates to 68° F.

300 LIFE HISTORY OF THE AUSTRALIAN OYSTER,

Churchill and Gutsell (1919) state that in Great South Bay, Long Island,
spawning began when the temperature of the water reached 70° F., and that it
proceeded rapidly while temperatures ranged from 70° to 74°, but that it slowed
down or ceased with temperatures of 70° to 68° F. When it rose to 74° F. the bulk
of the spawn was thrown out in the course of two or three days. In the same
locality Gutsell (1921) found that in 1921 spawning began with a water
temperature of 66:5° F. to 67° F.

Churchill (1920) states that O. virginica may spawn when the water reaches a
temperature of 68° F., but spawning proceeds at normal speed only when the water
is 70° or above.

T. C. Nelson (1921) found that, although spawning of oysters will commence
when the water has reached a temperature of about 70° F., it becomes much more
active with temperatures from 75° F. to 85° F. A rise in temperature is followed
by a more active spawning, while a decrease in temperature is accompanied by a
rapid falling off in the amount liberated.

Spawning was observed by Nelson on a float some six inches below the surface
of the water on 26th June, 1920, at 4.55 p.m., in bright sunlight. The tide was
about three-quarter flood, the density 1-018, and the temperature 78° F. Spawning
continued till 5 p.m. About one-third of the spawn was thrown out during this
spawning. An examination of the stomachs of the oysters showed that the oysters
had not been feeding during the duration of the spawning.

T. C. Nelson (1921a) reproduced a kymograph tracing made by two female
oysters which spawned while attached to an apparatus used to record feeding
reactions. The first oyster spawned on 6th August, 1920, from 4.18 p.m. to 4.29 p.m.
with the tide five-sixths flood, and the second on 26th July at 10.37 p.m. at high
tide.

Prytherch (1923) found that by placing oysters in tanks of slowly running
water, in the sunlight, spawning may be induced by the rising temperature. In
every such experiment from 10th July to ist September the oysters spawned
readily. Spawning occurred at temperatures ranging from 68° F. to 75° F., and
lasted over intervals of 15 to 30 minutes. At temperatures higher than 75° F., it
was found that the spawn was released slowly and immediately settled to the
bottom of the tank, whereas normally it would be forcibly ejected into the water
and would float about for some time. In one instance a group of 11 months old
oysters spawned for over five minutes at a temperature of 69:5° F. A sample of
this, observed in a watch-glass, developed as normally as that of the older oysters.

T. C. Nelson (1925) found a latent period of four days after the temperature
rose above 68° F. before spawning began.

The same author (1926) confirmed J. Nelson’s observations that after the
spawning temperature (68° F.) has been attained a sharp rise in temperature is
followed by spawning. He also found that spawning is most apt to occur between
late afternoon and midnight, and during the flood tide. T. C. Nelson observed or
recorded on a kymograph natural spawning of the oyster on five occasions. In
each instance the spawning took place in the daytime during the late flood tide
and with a rising temperature.

Galtsoff (1926) found that both male and female oysters may be induced to
spawn by increasing the temperature of the water, whereas spawning in the
female‘may be induced at a constant temperature by the addition of sperm. After
spawning, a female oyster is for two days immune to the presence of sperm,

BY T. C. ROUGHLEY. 301

T. C. Nelson (1927) observed that the more rapid the rise in temperature
above 68° F. the sooner spawning begins, though a higher temperature is required
to induce spawning than when the temperature rise has been more gradual and
extended over a longer time. In the latter respect Nelson’s observations on the
spawning of O. virginica are in entire agreement with those of Sparck (1925) on
O. edulis.

T. C. Nelson (1928a) stated that later spawnings occur usually at temperatures
above that which caused the initial spawning of the season, Following a cold
summer, unshed spawn may be found in the oysters even into the winter months.
The important observation was made, also, that the American oyster spawns after
the water temperature reaches 68° F. over all parts of its range, with no adjust-
ment to the extremes of its distribution.

Prytherch (1928) states that in many instances in 1925 and 1926 spawning
was observed to take place in tanks and floats (in Milford Harbour, Connecticut)
at temperatures ranging from 68° F. to 80° F. It was found that those oysters from
the Sound, where the water temperature is much lower than in the Harbour,
‘spawned in about half an hour at 68° F.-72° F., while the oysters from the warmer
harbour waters required several hours’ exposure to this temperature before
spawning occurred, or, on the other hand, they could be induced to spawn in half
an hour by increasing the temperature from 73° F. to 80° F.

During 1925 and 1926 the heaviest spawning of oysters in the harbour was
found to occur after the water had reached and maintained for a few days a
temperature of from 68° F. to 70° F. In both years the majority of the oysters
spawned at the end of the July full moon tidal period, when the temperature of the
water increased as the result of the greater range of tide during this period.
Prytherch maintains that the effect of the moon on the spawning of the American
oyster is only indirect, through the change it produces in the vertical and hori-
zontal movement of the water over the oyster beds leading to an increase of
temperature.

The discharge of spawn occurs near or at the time of high water. From
21st July to 29th July, 1926, the water temperature on the last of the ebb tide and
at low water was often from 68° F. to 79° F., and yet the oysters failed to spawn.
The same oysters, when placed in water pumped at high tide and warmed to the
same degree, spawned in a very short time. It was observed by Prytherch, also,
that oysters kept in floats always spawned near the time of high water, at
temperatures ranging from 68° F. to 75° F., and never at low water, though it was
several degrees warmer. In analysing the factors of temperature, salinity and
hydrogen-ion concentration at the time when spawning occurred, it was found that
the hydrogen-ion concentration or pH value of the water showed the greatest
difference. In all the observations but one, the water had a pH value ranging
from 7:8 to 8-2 when spawning occurred, while in the exceptional case the pH was
7-6 and the water temperature 73° F.

The failure of the oysters to spawn at low tide, when the temperature of the
water is often above 68° F. is thought by Prytherch to be due to the low alkalinity
of the water at this stage of the tide, as indicated by the pH readings.

Summarizing the studies of spawning in Milford Harbour, Prytherch con-
cludes that the most important controlling factors are the temperature of the
water, the range of the tide, and the hydrogen-ion concentration.

302 LIFE HISTORY OF THE AUSTRALIAN OYSTER,

T. C. Nelson (1929) found that oysters may spawn even though their gonads
may not be developed to their maximum extent; spawning may occur, in fact, when
the gonad is but poorly developed. He considers, however, that the crops of spat
resulting from such spawnings are very poor.

Galtsoff (1930a) found that the duration of the spawning period of O. virginica
varied from 36 to 70 minutes, and the number of contractions varied from 56 to
135, the average number of contractions per minute of four oysters being 1:36. In
the introduced Japanese species, O. gigas, he found that the duration of the
spawning period varied from 15 to 59 minutes, and the number of ‘contractions
from 31 to 121, the average number of contractions per minute of four oysters
being 1:96. The contractions of the two species are not strictly correlated, however,
for they were not observed at the same time and under identically similar
conditions, the record for O. virginica being obtained in July, 1929, and that for
O. gigas in October of the same year.

With regard to the larviparous oyster of the Pacific coast of the United
States (O. lurida), T. C. Nelson (1928) states that it spawns at the same tempera-
ture as O. edulis, viz., 59° F.—61° F. .

In the oviparous Japanese common oyster (O. gigas), Amemiya (1928)
observed numbers of oysters spawning both on the beds and in glass jars. On
the beds the oysters spawned about 30 to 40 minutes after they were submerged
(to a depth of about a foot) by the water on the flood tide. Unfortunately no
temperatures or other physical conditions are stated, and it is not recorded
whether the tide was spring or neap, though in another place Amemiya states
that the ordinary temperature which prevails in central Japan at the spawning
season is about 77° F. Amemiya observes that “Most probably in the natural
oyster bed, the impetus of the fresh current of the flood tide causes the stimulus to
spawning just as the change of water in the laboratory achieved the same result’.

Amemiya discusses the ejection of spawn by O. gigas when the water recedes
from the oysters on the ebb tide. He remarks that this snapping of the shells
on the ebb tide may be observed at times other than the spawning season of the
oyster, and he concludes that it is a “form of activity connected with some other
habits of the oyster, and is not employed solely for the purpose of spawning”’.
This snapping of the shells of oysters as the tide leaves them may be seen on
all oyster beds at all times of the year; it causes the ejection of the accumulations
of waste material rejected by the palps and carried along a ciliated track to the
mantle edge. By this means the mantle cavity is rid of all accumulated waste
prior to the shells being left exposed to the air, where they must remain without
a water circulation till the next tide covers them. It is in no way connected with
spawning.

Amemiya (1929) also states that O. gigas has one spawning season in a year,
and at the end of the season the animal is entirely exhausted.

With regard to the Japanese larviparous species, 0. denselamellosa, Seno
(1929) states that, in the Seto Inland Sea, spawning occurs most actively from
the end of June to the middle of July, when the water temperature ranges from
TOS UM, TKO) BOM,

In the case of the Bombay (oviparous) oyster (0. cucullata), Awati and
Rai (1931) found that, along the coast of Bombay, the optimum temperature for
spawning ranges from 80° F. to 87° F., and the optimum density is about 1-020.

BY T. C. ROUGHLEY. 303

On the coast of New South Wales spawning is a most irregular process.
With the warming of the water during the spring, the gonad develops rapidly
and may become fully developed by the middle of December. Spawning frequently
occurs in such oysters during the abnormally high spring tides round about the
Christmas period. These oysters frequently spawn again in late summer. If,
however, owing to an abnormally cool spring the gonad develops slowly, spawning
may occur in January, or even as late as April or May. In other cases there may
be several partial spawnings throughout the summer months. Spawning is not
confined to the summer months, however, for on rare occasions intermittent light
spawnings may occur right throughout the winter, as shown by an irregular
winter catch of spat. These have been noticed in the Hawkesbury River and at
Port Stephens. I have seen, in November, young spat attached to the mangrove
sticks from the size of about a millimetre ranging up to about 10 millimetres,
which must have developed from winter spawn. When, as is usual, spawning is
completed by the end of summer, the gonad passes through a quiescent period
during the winter as far as the development of ova or sperms is concerned, but
activity is displayed in the storage of glycogen in the vesicular tissue in prepara-
tion for the development of ova and sperms in the spring. The formation and
storage of glycogen, however, does not appear to be so heavy as in the European
oyster (0. edulis), which is frequently spoken of as fattening during the winter
months. The size of the gonad of the New South Wales oyster in the winter is
not comparable with its size in the summer, when it is distended with spawn. Nor
is it appreciated to the same extent as food by the public, who prefer it with a
well developed gonad. The winter in New South Wales is a period of greatest
shell growth in preparation for the active development of the gonad during the
spring and summer.

The irregularity of the spawning period in this species is no doubt largely
accounted for by the fact that the great bulk of the oysters in New South Wales
are grown in the tidal zone where temperature fluctuations, varying from
cold water to hot sun in the course of a few hours, are enormous.

The vertical range of the New South Wales oyster extends from about mean
high water level to a depth of upwards of 20 metres, and prior to the seventies
of last century most of the oysters marketed were taken from permanently
submerged beds. About 1870, however, the activity of a small worm (Polydora
ciliata), which appears to have first made its appearance in the Hunter River,
began to kill off vast numbers of these oysters. Gradually the worm spread
to other rivers, and at the present day it is impossible to raise oysters on
submerged beds in most of the rivers of New South Wales. The Manning and
Bellinger Rivers are, however, notable exceptions; in the former the worm is
present, but does very little damage, and in the latter it does not appear to
occur at all. The mud worm has therefore forced cultivation to the tidal zone,
where it can be combated.

If the oysters in New South Wales were grown on permanently submerged
beds, where the temperature is not subject to the violent fluctuations characteristic
of the tidal zone, it is reasonable to expect that spawning would be more regular
in its occurrence, and an indication of an impending spawning given by the
gradual rise in the temperature of the water, as obtains in America, where the
bulk of the oysters are grown in situations never bared by the tide.

304 LIFE HISTORY OF THE AUSTRALIAN OYSTER,

Experiments in Artificial Fertilization.

Experiments were carried out at various periods in order to study the
embryonic development of the oyster, the rate of growth, and the influence of
temperature on the growth-rate, and to compare the rate of development with
that of other oysters, particularly O. virginica of the Atlantic coast of America.

At Port Macquarie in 1924 artificial fertilization experiments were carried
out in water of four different densities. In A, ordinary estuarine water with a
density of 1:021 was used; in B, tank water was added in sufficient quantity to
lower the density to 1:015; in C, to 1:011; and in D, to 1:005. The initial
temperature of the water in each case was 74° F. It was found that the rate of
development decreased as the density was lowered. The free-swimming stage was
reached in A in 8 hours; in B in 9 hours; and in C in 10% hours. No free-
swimming embryos at all developed in D, and an examination 21 hours after
the mixing of the eggs and sperms showed that fully 50% of the eges had
remained unfertilized, and of those in which fertilization had been effected,
little development had taken place. There was no development beyond the
morula stage.

The embryos in A, B, and C were kept under observation for 70 hours, and
in each case appeared to develop quite normally. The temperature of the
water varied during the course of the experiment from 70° F. to 74° F.

While the oysters were spawning on a bed at Port Macquarie on 5th April,
1924, a large accumulator jar was filled to a depth of 8 inches with water over
the spawning bed. The temperature of the water was 70° F., and its density
1:021. In this water four spawning oysters were placed and allowed to remain
till 10.30 p.m. on the same day. In a short time the bottom of the jar was
white with spawn. At 9 p.m., seven hours after fertilization, a small percentage
of the embryos was found to be swimming on or in the vicinity of the bottom.
The temperature of the water had fallen in the meantime to 68° F. At 10 p.m.,
eight hours after fertilization, numbers of embryos were found at the surface of
the water, having swum vertically a distance of eight inches. A quarter of
an hour later numerous short, white columns were seen extending downwards
from the surface of the water; these were formed by congregations of embryos
swimming towards the bottom. These columns are characteristic of the free-
swimming, embryonic stage of the oyster. The embryos are continually
swimming from the bottom to the surface, where they may remain for some
time attached to the surface film, before making their way again to the bottom.
They rise singly, and in their passage cannot be detected by the naked eye, but
in their journey downwards they follow each other in well-defined lines. The
columns are distributed right throughout the water, but are usually more
prevalent near the sides of the jar. Vibration of the containing vessel, which
causes a slight agitation of the water, stimulates the embryos at the surface
to descend. The columns are most numerous near the surface of the water,
but nevertheless many of them extend right to the bottom @2L yeah, ile Bil) 2

After about 9 hours the great bulk of the embryos were freely swimming in
the jar, and the bottom was practically clear.

At 12, midnight, on the day following the spawning, 34 hours after fertilization,
the first embryos were seen completely covered with shells, the hinge-line being
straight. The temperature during this time had varied from 68° F. to 70° F. The

BY T. C. ROUGHLEY. 305

average measurements of 20 larvae at this time were: length, 67; depth, 53; and
the length of the hinge, 45y.

It is characteristic of the larvae to swim in circles, and when moving from
the bottom to the surface of the water, their course is therefore a long spiral
of narrow diameter. Shelled larvae, on reaching the surface, may remain there
for some considerable time with the valves of the shells extended, the ciliated
velum attached to the surface film. A sudden withdrawal of the velum and
snapping together of the valves causes the larva to sink to the bottom, where,
in a very short time, it will open its shell, protrude the velum, and again swim
spirally to the surface. Larval life, then, consists of voluntary movements
up and down in the water, the larvae at the same time being carried hither and
thither by tides and currents.

After three or four days the larvae, which till then were practically devoid of
colour, developed a bluish or bluish-grey tint, most pronounced dorsally in the
vicinity of the hinge.

At the end of 5 days the average dimensions of the larvae were as follows:
length, 754; depth, 584; and the length of the hinge, 50u (PI. xxv, fig. 32). No
special effort was made to carry the larvae through the whole of their free-
swimming period, and on the sixth day the prolific development of infusoria
in the water rendered it necessary to discontinue the experiment.

On a number of occasions the rate at which the embryos swam was measured
with the aid of an ocular micrometer. A drop of water containing embryos was
placed on a microscopic slide and a cover-glass gently lowered over it. By means of
a stop-watch the time taken by the embryos to traverse the full length of the
micrometer scale was ascertained, and an average obtained from numerous counts.
The total length of the micrometer scale was 1/31 inch. On one occasion, embryos
69 hours old, which had not developed shells, took an average of 234 seconds to
swim the length of the scale; at the rate, in other words, of approximately 46
inches per hour. On another occasion, embryos 24 hours old were timed to swim
at an average rate of approximately 39 inches per hour, and at the end of 30 hours
at about 58 inches per hour. The rate of swimming of different batches at
different ages, therefore, ranged from 39 inches to 58 inches per hour. In view,
however, of the artificial conditions under which these embryos were living,
lacking efficient aeration, etc., it cannot be assumed that their progress under
natural conditions would be the same; their vitality and therefore their activity
would in all probability be greater. The rate given here may be taken as a
conservative estimate of the progress of the embryos under natural conditions.

In order to determine the part played by the temperature on the rate of
development of the embryos of O. commercialis, the following experiments were
carried out.

Eggs and sperms were mixed in water obtained from the Hawkesbury River
in two large accumulator jars, the depth of water in each case being 10 inches
and the density 1-016. In one case, A, the initial temperature of the water was
77° F., and in the other, B, 64° F.

In A, at the end of 14 hours many ova had completed their first division;
after 12 hours numbers of ova showed a partial division into two micromeres
and a deutomere; after two hours many showed four micromeres; after 23 hours
the early morula stage was seen; after 5 hours the embryos, now ovate in form,
had reached the gastrula stage. After 6 hours several embryos had reached the

306 LIFE HISTORY OF THE AUSTRALIAN OYSTER,

swimming (trochophore) stage, revolving in circles, the cilia beating so rapidly
that, even with the critical definition of an oil lamp, they could not be dis-
tinguished. All embryos, when beginning to swim, revolve in small circles on the
bottom, mostly, but not always, in a counter-clockwise direction. After 6% hours
several embryos, when transferred from the accumulator jar to a watch-glass,
were seen to leave the bottom and swim upwards for short distances. After 9
hours several embryos were found at the surface of the water in the accumulator
jar, ten inches from the bottom.

During the course of this experiment, the temperature, which at the beginning
was 77° F., varied from 76° F. to 80° F. The temperature was taken every
half-hour and the mean determined as 78° F.

In experiment B, the temperature of the water when the eggs and sperms
were mixed was 64° F., the depth of the water (10 inches) in the accumulator
jar, and its density, 1:016, being the same as in experiment A.

At the end of an hour, some fertilized ova had become spherical and more
opaque; the majority, however, were still more or less pear-shaped, though they
had developed an opacity indicative of fertilization.

After 2 hours partial cleavages only were seen. After 23 hours several embryos
were seen with a division into two micromeres and a deutomere. After 34 hours
a number were seen with four micromeres; and after 5 hours the most pronounced
development noted was an early morula. (It will be noted that this stage was
reached in 23 hours in experiment A.) After 11 hours many embryos had reached
the early gastrula stage; at the end of 22 hours the first swimming embryos were
seen revolving very slowly.

During this experiment the temperature varied between 55° F. and 65° F.
The experiment was begun at 1.15 p.m., and from then till midnight the temper-
ature fluctuated between 64° F. and 60° F. At 7 a.m. the following morning it
had fallen to 55° F., whence it gradually increased till at 11.15 a.m. it had risen
to 59° F. From midnight till 7 a.m. the following morning, no temperatures were
noted; with the exception of this period the temperatures were taken half-hourly,
the average working out at 62° F. Owing to a decided fall during the intervening
blank period, the average for the whole period would be somewhat lower than this.

A comparison of the rate of development of the embryos in each experiment
shows a marked disparity due to the difference in the temperatures. In the first
case, at an estimated average temperature of 78° F., the free-swimming stage was
reached in 6 hours, while in the second, at an estimated average temperature of
62° F., the time taken to reach the same stage of development was 22 hours.

An average temperature of 78° F. is somewhat higher than that which would
obtain in the estuarine waters of New South Wales during the summer months,
and therefore it may be presumed that the time taken by the oyster embryos to
reach the free-swimming stage will be somewhat longer than 6 hours. The
estimated average temperature which obtained during the course of the second
experiment, 62° F., is considerably higher than that which prevails during the
late winter months. During an investigation of a winter mortality of oysters in the
George’s River in 1924 (Roughley, 1926), I found that at a depth of 2 feet 6 inches
below mean low water level the temperature varied, during July and August,
from 50° F. to 59° F. The highest temperature reached was lower than the
average of this experiment. During the winter, therefore, embryonic development
will probably be somewhat slower than that indicated during the course of this

BY T. C. ROUGHLEY. 307

experiment. Of course, oyster embryos have rarely to face a winter temperature,
but, as we have seen, winter spawnings do occasionally occur.

An effort was made both at Port Macquarie during 1924 and at the Hawkesbury
River during 1925 to determine the duration of the free-swimming stage, following
the method initiated by Stafford in Canada in 1907, which consisted of collecting
the larvae in a plankton net daily, and making accurate measurements of the
predominating size-group and recording the daily increase in growth.

During my six weeks’ stay at Port Macquarie during the late summer of 1924,
the plankton net was towed at least once a day, in various situations in the
estuary, but on not one occasion were any oyster larvae found. After the spawning
which occurred on the beds near the boatshed where my laboratory was situated,
the net was used with greater frequency, at various states of the tide and in
many and varied situations, both at the surface and close to the bottom in shallow
water, but on no occasion were any larvae recovered. After the lapse of a fortnight
the attempt was abandoned, but not before I felt justified in predicting that, in
spite of the spawning, a set of spat was unlikely to occur. And so it turned out,
though I did not receive any indication of the conditions which operated to destroy
the larvae.

Again, at the Hawkesbury River during 1925, I used the plankton net at
least once daily, not only near the State Fisheries boatshed at Brooklyn, which
was fitted as a temporary laboratory, but in situations as far removed as Cowan
Creek, near the entrance of the river into Broken Bay, and also many miles
upstream. On every occasion oyster larvae were obtained, both in the straight-
hinge and umbo stages, but never was there a definite preponderance of an age-
group (Pl. xxv, fig. 33). This was rather surprising in view of the fact that
examinations of oysters’ gonads on many occasions previously had indicated that
spawning occurred mostly during spring tides of either full or new moons. It
became apparent that in the Hawkesbury River, during the period under review,
the spawning was either not confined to spring tides, or the scattering of the
larvae subjected them to a range of temperature sufficiently wide to cause a
marked disparity in the growth-rate. Careful analysis of each day’s plankton
collection gave the impression that the spawning of the oysters in the Hawkesbury
River during that period was occurring almost daily. The beds are distributed
over a very great area, extending from the entrance to a distance of about 20
miles upstream, and also in numerous large creeks which bear heavy crops of
oysters, extending in each case for many miles. Spread over such an area, a wide
range of conditions is met with, both as regards salinity and temperature, and a
uniform spawning period can scarcely be expected. On the contrary, one would
anticipate great irregularity; and it is probable that under such conditions
spawning occurs on different beds at different times throughout the summer
months.

Most crops of very young spat which I have examined have shown a similar
diversity of age, but on one or two occasions I have seen a general uniformity in
the size of very young attached spat. In order to determine the length of the
larval life of the oyster in New South Wales, therefore, one would have to be
fortunate enough to carry out an investigation when the spawning and growth of
the larvae showed a pronounced degree of uniformity.

In the absence of direct evidence of the length of larval life of the Australian
oyster, an indication of its duration may be obtained by a comparison of the

308 LIFE HISTORY OF THE AUSTRALIAN OYSTER,

growth-rate during the embryonic stages with that of the oyster of the Atlantic
coast of America (O. virginica). At a temperature of about 80° F. the free-
swimming stage in O. virginica is reached, according to T. C. Nelson (1921), in
from four to five hours; and we have seen that, at a temperature of about 78° F.,
the same stage is reached in O. commercialis in six hours. Nelson also states
(loc. cit.) that the larva is completely enveloped in shells within 24 to 36
hours after fertilization, while the Australian oyster took 34 hours to reach that
stage at a temperature varying from 68° F. to 70° F. A difference in temperature
of 2° as recorded in the American and our experiments would cause but a slight
difference in the rate of growth, and it would appear, therefore, that the Australian
oyster develops somewhat more slowly than the American oyster at the same
temperature. If the subsequent growth of the larvae is maintained at the same
relative rate, it is apparent that the free-swimming life of the Australian oyster
will be somewhat longer than 14-16 days.

The absence of predominating size-groups in the Australian plankton and
their more regular occurrence in the American plankton may be explained by
the differences in the habitats of the parent oysters. In Australia the great bulk
of the oysters occur in the tidal zone, where they are subject to great variations
of temperature, leading to an irregularity of spawning; in America, however,
most of the oysters are never exposed by the tide, often growing in water of
considerable depth where there is a certain uniformity of temperature, which
must tend to a greater uniformity of sexual development and a more regular
spawning.

In the United States of America the determination of the occurrence of oyster
larvae in the plankton is a question of considerable economic importance, as has
been demonstrated by T. C. Nelson on a number of occasions. Oyster cultivation
in America consists, in the main, of dumping large quantities of shells on the
bottom to provide attachment surfaces for the larvae. Now, the efficiency of any
form of cultch is dependent very largely on its cleanliness, and its cleanliness is
governed by the length of time it remains in the water. Material which has been
submerged for a month or two will not catch nearly so many oysters as that which
has been in the water for a few days only, owing to the deposition of silt over
the surface. If, then, an examination of the plankton reveals large quantities of
oyster larvae of approximately the same age, it can at once be determined when
the subsequent fall of spat may be expected, and the shells can be dumped on
the bottom within a few days of the spatting. Thus the largest possible crop is
obtained.

A knowledge of the length of larval life does not promise to be of similar
importance to the oyster fisheries of Australia, at least so long as the bulk of the
oysters are grown in the tidal zone, for spawning is always likely to be inter-
mittent, and this naturally leads to an irregular catch of spat, extending over a
considerable period.

Orton (1926), in O. edulis, determined that the larvae under natural
conditions are retained in the mantle cavity of the parent for a period varying
from 1 to 14 weeks, and that, generally, settled spat may be seen in fair numbers
in normally warm weather about a month after the first batch of white-sick
oysters is seen.

Hori and Kusakabe (1927) succeeded in raising the larvae of the oviparous
O. gigas to the setting stage in petrie dishes by feeding them on a minute alga

BY T. C. ROUGHLEY. 309

(Chlorella pacifica). Under these conditions the length of larval life was 23 days,
at a temperature which fluctuated between 73° F. and 90° F., the average at 10 a.m.
being 82° F. The density of the water was about 1:019-1:021. It is reasonable to
presume, however, that this rate of development will be found to be considerably
slower than that which obtains under natural conditions.

In the Japanese larviparous oyster (0. denselamellosa), Seno (1929) states
that fixation takes place about 4 weeks after spawning, at an approximate water
temperature of 68° F.

Further observations on the development of oysters of various species may be
noted for comparison with that of O. commercidlis.

In_O. edulis, Huxley (1883) states that the larva at the time of ejection from
the parent measures 1/150 inch (0-17 mm.) long. In the larviparous Australian
oyster (O. angasi), I have found shelled larvae as large as 0:2 mm. The largest
shelled larvae found by Stafford (1913) in the mantle cavity of the oyster of the
Pacific coast of America (O. lwrida) measured 0-186 mm.

In O. edulis, Sparck (1925) states that the larvae cannot complete their
development in water with a salinity below 24-25%,. In the same species, Orton
(1926) found that the larvae develop to the shelled stage in about a week at
summer temperatures.

Orton (1927) also states that artificial fertilization cannot be performed at
present in the case of O. edulis with much chance of obtaining a normal rate of
development, for when embryos are taken away from the parent, and are left in
unchanged ordinary sea water, development soon becomes abnormal.

In O. virginica, T. C. Nelson (1921) states that when first completely covered
with shells the larvae measure 0:06 mm. in length. Prytherch (1923) found that at
a temperature of 80°-85° F. the larvae invariably died; the most successful
development occurred at a temperature of 70° F. T. C. Nelson (1921) had already
noted that the embryos begin to swim in from 4 to 5 hours after fertilization at
a temperature of 80° F.

In O. gigas, Hori (1926) found that the larva forms a bivalve shell in from
2 to 3 days; its dimensions at this stage are: length, 0:07 mm.; height, 0-06 mm.;
and length of hinge, 0:04 mm. Seno, Hori and Kusakabe (1926) determined that the
optimum temperature for larval development of O. gigas lies between 73° F. and
79° F., and the optimum density between 1-017 and 1:021. Amemiya (1928) states
that the larval shells of O. gigas grow large enough to cover the whole body and
velum in from 8 to 4 days. The time required for the development of
O. denselamellosa is stated by Amemiya to be much longer than that of other
Japanese oysters.

The Sex Ratio of O. commercialis.

During the past 15 years I have on many occasions examined microscopically
the gonads of this species, and whenever a batch was worked through I have
invariably noticed that the females predominated. In order to determine the ratio
of females to males over as much of the State as possible at approximately the
same time of the year, I had forwarded to me during January and February, 1927,
through the courtesy of State Fisheries, 100 oysters from the most important
oyster producing rivers in the State, numbering 27 in all. From three of these
rivers two consignments were received, so that a total of 3,000 oysters was
examined. The proportion of females to males is shown in the following table:

(0)

310

LIFE HISTORY OF THE AUSTRALIAN OYSTER,

Locality fe) 3 Locality. Q 3
Tweed River 64 36 Brisbane Water 68 32
Brunswick River 62 38 Hawkesbury River 70 30
Richmond River .. 74 26 George’s River 85 15
Evans River 82 18 Crookhaven River 78 22
Clarence River 81 19 Currumbene Creek 70 30
Sandon River 72 28 Clyde River 68 32
Bellinger River 66 34 Moruya River 78 22
Nambucca River 82 18 Wagonga River 68 32
Port Macquarie 82 18 Wallaga Lake .. 70 30
Manning River 75 25 Wapengo River 58 42
Wallis Lake wo 70 30 Nelson Lake 84 16
Lower Port Stephens .. Oe 54 46 Merimbula Lake an ae 64 36
Upper Port Stephens .. Pre 84 16 Pambula River ae ue 76 24
Hunter River sé ae . 88 12

It will be noted that in every case the females predominated, ranging from 54%
in Lower Port Stephens to 88% in the Hunter River. The average percentage was:
females 73%; males 27%. In other words, there were 2:7 times as many females
as males. The highest percentage of females I have ever found occurred in a batch
of 100 large oysters from the Hawkesbury River, which were examined on 31st
March, 1931. The females numbered 96%.

Toward the end of this investigation I opened 15 very young oysters which
were attached to the larger ones received from Brisbane Water, and noticed that
the gonads were just showing signs of development, the ducts being in most cases
just visible. Every one was found to be a male, containing actively moving sperms,
but no ova in any stage of development.

In a further batch received from Brisbane Water there were 8 very young
oysters, and these, too, proved to be all males. In the consignment from the
Clarence River, also, there were 9 very young oysters attached to the older ones;
again, all were males. Amongst those received from Lower Port Stephens were
19 very young individuals, all of which were males. Similarly, 16 young oysters
from the George’s River, and 10 from the Manning River were all found to be
males.

A sex-change during the course of development was at once indicated, and
steps were immediately taken to procure for examination further samples of very
young oysters. A hundred young oysters were received from each of the following
localities: Port Macquarie, Lower Port Stephens, Pambula River and the Hawkes-
bury River.

The oysters from Port Macquarie were received on 1st February, 1927, and it
was stated by the Inspector of Fisheries who forwarded them that they were from
a crop that had set during March the previous year. This would give them a
maximum age of 10 months. The gonads were very poorly developed, the digestive
diverticula in the bulk of them not having yet been obscured. It was found that
84% were males and 16% females.

The oysters from Port Stephens had caught on mangrove sticks which had
been laid out in January, 1926, and were reputed by the Inspector of Fisheries to
have attached in the following month. They were examined at the end of February,
1927, and their greatest age could not, therefore, have been more than 13 months.

BY T. C. ROUGHLEY. 311

There was, however, no uniformity in the size or age of the oysters, several crops
of various ages being indicated. It was found that 66% were males, 19% females,
18% contained neither ova nor sperms, and 2% were hermaphrodite, containing
well developed ova and motile sperms. Some of the older oysters had well
developed gonads, and amongst these the females occurred in greater numbers
than amongst the smaller individuals, which were almost invariably males. In
two instances, however, very young oysters showed developing ova.

The oysters received from the Pambula River were generally larger than those
from Port Stephens, and the gonad was much better developed, some showing very
pronounced development. They varied greatly in size, however, and although they
were described by the Inspector of Fisheries as being a year old, there was
clearly considerable variation in the ages of different individuals. It was found
that 65% were males; 28% were females; and 3% were hermaphrodite, containing
ova and sperms in various stages of development.

It will be seen that in three samples of oysters from different rivers, the oldest
of which were reputed to be not older than 14 months, the males considerably
outnumbered the females. With very few exceptions, the youngest oysters, in
which the gonad was in an early stage of development, were males. It is quite
apparent, therefore, that the Australian oyster (O. commercialis) undergoes a sex-
change, but it is not yet quite clear whether it always functions first as a male.
It certainly does in the great majority of instances, but a few individuals have
been examined with the gonad in a very early stage of development, in which
ova and not sperms were developing. It is possible, of course, that these oysters
were not exceptions, that they had already functioned as males, though of this
no indication could be obtained. It may be regarded as significant, however, that
in the first instance, a total of 77 very young oysters received from five different
rivers proved, without exception, to be males, and the examination of further
batches during the following month showed a small percentage functioning as
females. Is it possible that some of the oysters had spawned as males in the
meantime, and had then begun to develop ova?

When this indication was given that a sex-change occurred, a careful
examination was made of each individual in an effort to locate both ova and sperms
in the one oyster. Previously, microscopical examination had been confined to the
use of a 2/3 inch objective with ordinary illumination, which was quite service-
able for detecting ova or sperms when the field was confined to either one or the
other. If, however, both ova and sperms occurred in the same field, there was a
risk of overlooking the sperms, which are relatively very small, or, perhaps, con-
fusing them with the Brownian movement of particles of protoplasm from damaged
ova. For this reason, then, every smear from the gonad was subsequently examined
first with a 2/3 inch objective illuminated through an ordinary condenser, and
then with a 1/10 inch objective using a dark-ground condenser, with which sperms
could be detected with unmistakable clearness. Later, on account of the time
occupied in the use of the dark-ground condenser, the examination for sperms
was made by dark-ground illumination using a 1/3 inch objective and an expanding
stop, by means of which a sufficiently large and perfectly clear image of sperms
was obtained. By this means a number of oysters was found with both ova and
sperms in the gonad. Mention has already been made of the fact that an
occasional young oyster contained both ova and sperms in the gonad, but it was
found that this condition is not confined to the young stages, for, amongst 100
marketable oysters (approximately three years old) received from the Tweed

312 LIFE HISTORY OF THE AUSTRALIAN OYSTER,

River, two contained both ova and sperms. In one both were in abundance, and
in the other there were many sperms but relatively few ova. One individual was
also received from the Hawkesbury River which contained both ova and sperms in
abundance. A total of nine hermaphrodite oysters was found. It is possible that
other examples of hermaphrodite oysters would have been found if both ova and
sperms had been carefully searched for during the whole of the examination of
the three thousand oysters previously referred to; it was unfortunately late in
the examination that an indication of a sex-change was revealed, and therefore a
small proportion only of the three thousand oysters was examined critically with
this object in view.

Microscopic sections were prepared of three young oysters whose approximate
age was twelve months, and of two large marketable oysters which I should
estimate to be about three years old.

In an oyster taken from the Pambula River on 1st March, 1927, reputed to be
about twelve months old, the shell of which measured 14 inches in length, most of
the germinal ducts contained both ova and sperms (PI. xxii, fig. 26, o., s.), the
sperms filling up the lumen of the duct, while the eggs were in various stages of
development from the germinal epithelium. A few of the ducts contained eggs
only, and in a few restricted areas the ducts were filled with well developed ova
while spermatocytes were developing from the germinal epithelium. The follicles
varied greatly in depth; in some situations where the vesicular connective tissue
was narrow they extended as far as the digestive diverticula, in others, where
the depth of the vesicular tissue was much greater the follicles extended only
about half way. Most of the follicles were of small diameter; in most situations
the wall of the follicle had attached to it eggs in various stages of development,
and enclosed an abundance of ripe spermatozoa; in others the follicle enclosed
large, well-developed eggs with spermatozoa in the spaces between them (Pl. xxii,
1s, PANS Don Os S5))o

In this oyster, therefore, sperm development appears for the most part to
have preceded egg development, while in restricted areas both eggs and sperms
continued to develop side by side.

In an oyster taken from the Hawkesbury River on 3rd February, 1927, which
measured 1% inches long, the gonad was much better developed than that just
described, and contained eggs and sperms in about equal proportions. For the
most part developing eggs were attached to the walls of both the ducts and
follicles, and surrounded masses of ripe spermatozoa. It is clear that the bulk of
the sperms have developed prior to the development of the eggs, but in some
situations developing eggs and spermatocyte adjoin each other in the same duct
or follicle, in some cases with eggs, and in others with spermatocytes, predomin-
ating. Active egg and sperm development is therefore taking place concurrently.

In a twelve months old oyster taken from Port Stephens on 28th February,
1927, the shell of which measured 1% inches, the gonad was but poorly developed,
the ducts just becoming apparent at the surface. In this oyster eggs were found
to predominate, and many of them appeared to be fully developed. Many of the
ducts were filled with eggs only, while eggs only were developing from the
germinal epithelium; in others the bulk of the lumen was filled with eggs with
sperms clustered in the spaces between them, sometimes with a dense mass lying
against the ciliated epithelium. In a few of the ducts and follicles the development
of both eggs and sperms was very active. From the advanced stage of egg
development in this oyster, it is apparent that the proliferation of both eges and
sperms has gone on side by side for a considerable time.

i)

BY T. C. ROUGHLEY. 31é

In a marketable oyster taken from the Tweed River on 22nd February, 1927,
in which the gonad was very well developed, microscopic examination of sections
of the whole of the gonad showed that on the right side an extensive sperm
development had been followed by very active egg development; in one or two of
the ducts eggs predominated, but all contained spermatozoa in contact with the
ciliated epithelium, while deeper down, the follicles, which were for the most part
small and separated by considerable amounts of unabsorbed vesicular tissue, were
all bordered by developing ova and enclosed compact masses of sperms (PI. xxili,

g. 28). In some situations both eggs and spermatocytes were actively developing
in the same follicle. Dorsally, between the origins of the mantles, this condition
gave way to a preponderating sperm development, whilst on the left side the gonad
almost entirely consisted of testis; the ducts were filled with spermatozoa, and the
follicles lined by spermatogonia and spermatocytes, which enclosed large masses
of sperms. In a few isolated situations, however, a few eggs were found attached
to the follicular wall, while deep down in the testicular tissue an occasional
follicle was seen which contained ova only in various stages of development. It
is apparent, therefore, that, although great numbers of spermatozoa were produced
before a general tendency occurred to produce eggs, a few eggs were differentiated
very early in the sexual development of the oyster during that season. If, when
this oyster was alive, some of the sexual products of the gonad on the left side
were examined, it is probable that sperms only would have been found, and the
oyster would therefore have been identified as a male; if, however, the sexual
products were obtained from the right side, an abundance of both eggs and sperms
would have been found, and the true. hermaphrodite condition of the gonad
disclosed.

In another marketable oyster from the Tweed River which occurred in the
same batch as that just described, the gonad was well developed, the areas of
unabsorbed vesicular tissue being very small and inconspicuous. Great numbers
of eggs and sperms occurred in about equal bulk throughout the whole of the
gonad. The ducts, most of which had coalesced with their neighbours, contained
for the most part an abundance of spermatozoa in contact with the ciliated
epithelium; in some cases, however, eggs lay against the cilia; and in others again
the eggs and sperms were intermingled indiscriminately. The convoluted follicles
(Pl. xxiii, fig. 29) were lined by eggs in active development, all stages from the
smallest to the mature egg being abundant; they enclosed irregular masses of
sperms. Sperm development, however, had almost entirely ceased, for very few
spermatocytes could be found.

Observations on the sex-ratio of other species of oysters have been made by
a number of authors.

In the European larviparous oyster, Dantan (1913) found that amongst one-
year old oysters 76% were males, and 6% females; when two years old 84% were
males, and 15% females; and when three years old 81% were males, and 18%
females. Dantan also found that the largest oysters always contained spermatozoa,
and that eggs are only found in those of medium size. He thought it likely that
the placing of individuals under less favourable conditions favoured the formation
of maie elements.

Sparck (1925) found that in the years since 1919 there were at no time in the
Limfjord more than 8%, to 10% of females; while from 75% to 80% of individuals
were found containing spermatozoa. In the colder waters of the Limfjord females
are found principally amongst the older oysters.

314 LIFE HISTORY OF THE AUSTRALIAN OYSTER,

Orton and Awati (1926) found that females predominated in the oviparous
Portuguese oyster, O. (Gryphaea) angulata, which had been relaid on English
oyster beds, the proportion varying from 60% to 88%.

Stafford (1913) states that the sexes of the American oyster (0. virginica)
are approximately equal in numbers. Similar results were obtained by T. C. Nelson
and other American authors.

In the Japanese oviparous oyster (0. gigas), Amemiya (1928) found that the
sex-ratio varied with the amount of nourishment obtained. On a good fattening
ground the ratio of females to males was 100: 95 amongst the one-winter group,
and 100: 73 amongst the two-winter group. On a poor fattening ground, the ratio
of females to males was 100: 116 amongst the one-winter group, and 100: 155
amongst the two-winter group. Amemiya concludes, therefore, that an abundance
of food tends towards the development of a preponderance of females, and a paucity
of food tends towards the development of males, thus to some extent bearing out
Dantan’s conclusion in the case of O. edulis.

In O. mordar, an oyster which occurs prolifically on the Great Barrier Reef
and North Queensland coast, an examination of 100 oysters from Hthel Rocks,
near Gladstone, on 9th March, 1931, yielded more than twice as many males as
females, the percentages being: females, 29%; males, 65%; indeterminate sex,
6%. In a communication to me from the Marine Biological Station, Low Island,
North Queensland, a little later, F. W. Moorhouse stated, however, that an
examination of oysters of this species made by him at Low Island yielded 62%
females; and 37% females; with 1% of undetermined sex.

A brief outline of our knowledge of the sex-change in various species of oysters
has been given in the introduction. Following the more detailed discussion of the
sex-change in the Australian O. commercialis, further observations on its occur-
rence in other species will serve as an interesting comparison.

Davaine (1852) and Hoek (1883) stated that a sex-change occurs in O. edulis,
and they based their conclusions on the occurrence of developing spermatozoa in
individuals which had recently functioned as females. It has long been established
that spermatozoa develop in oysters which still retain embryos or larvae in the
mantle cavity, and which, therefore, had recently spawned as females, but it
remained for Sparck (1925) to produce direct evidence of the change from male
to female. By boring through the shells of male oysters, and re-boring them later,
it was found that the spermatozoa had given place to eggs. Sparck is further of
the opinion that external conditions play a big part in regulating the sex, and an
alteration in the surrounding conditions results in an alteration of the kind of sex.

In the Portuguese oyster, O. (Gryphaea) angulata, Amemiya (1925-6) found
that after 75 individuals had been confined to a tank for a month, two had
developed a hermaphrodite condition. He is of the opinion that in the oyster the
genetic sex-determining mechanism, even though it may incline the future
differentiation of the individual towards either the male or the female type of
sexual organization, is not all-powerful; the sex of an individual is not irrevocably
determined by this mechanism. In the two cases of hermaphroditism referred to
above, the earlier phases of their gonadic differentiation were pursued under
conditions of ample nutrition: such of the gonadic tissues as differentiated at this
time therefore became ovarian tissues. When placed in the tank, however, where
food was relatively scanty, the gonadic tissues which developed under these
adverse conditions assumed the organization of testis. In other words, an abund-

ol

BY T. C. ROUGHILEY. 31

ance of food leads to the development of the physiologically more expensive
ovarian tissue, and a scarcity will tend towards the development of testicular
tissue.

Amemiya (1929) examined the gonads of one-winter individuals of 0. gigas by
boring through the shells, and repeated the operation when they had passed the
second winter. Two batches were examined. In the first batch, examined in 1928
(one-winter oysters), 119 were females and 58 were males. In 1929, the original 119
females were found to consist of 82 females and 20 males; and of the 58 males, 32
were found to be females, and 18 were males. In the second batch examined in
1928 (one-winter oysters), 209 were females and 145 were males. In 1929, the
original 209 females consisted of 85 females and 34 males; and of the 145 males,
48 were females, and 39 were males. In the first batch 29% had changed sex, and
in the second, 23%. The sex-change amongst the total oysters examined was 25%.

Amemiya considers it probable that at the beginning of every new spawning
season, the sex differentiates independently of the sex of the preceding season, and
that protandry does not occur.

In the Bombay oyster (0. cucullata), Awati and Rai (1931) found that
amongst normally healthy individuals 56% were females, and 41% were males,
3% being hermaphrodite, while amongst those which were harbouring pea-crabs
(Pinnotheres) in the mantle cavity, only 10% were females, and 83% were males,
7% being hermaphrodite. It is thought that the pea-crab influences the change of
sex either by reducing the food supply of the oyster, or by bringing about a change
in its general metabolism.

In view of the observations of the above-mentioned authors on the bearing
which the abundance or paucity of food has on the sex-determination, I obtained
several batches of oysters from various localities in an endeavour to discover
whether the food supply has any influence on sex differentiation in O. commercidlis.

On 17th March, 1931, I examined two batches of oysters from the Clyde River,
one from a bed about 13 miles from the entrance, described by the Inspector of
Fisheries as the finest growing and fattening ground in the river; the other from
outside the bar entrance, where the salinity of the water is always high, and
growth is very slow. In the first batch the females numbered 66%; the males 33%;
and one was hermaphrodite; in the second batch the females were 65%, and the
males 35%. It may be presumed that food is much more abundant on the bed
where growth is very rapid than in the locality where growth is very slow, yet
the percentage of females was practically the same.

Again, on 25th March, 1931, I examined a batch of oysters from Burraneer
Point, Port Hacking, where conditions are very unfavourable to oyster life. These
oysters, although in most cases of considerable age, were small and stunted, with
very hard shells. They were obviously growing where food is never abundant, yet
65% of them were females, and 35% males.

I have a further record of the sex of 100 oysters from Middle Harbour, Port
Jackson, examined in 1922. Here, again, conditions are very much against rapid
development, and the oysters rarely reach a marketable size, yet 64% of these
were females, and 36% males.

Here are records, then, of three batches of oysters from areas where the
conditions are very adverse for oyster development, and presumably where the
food supply is poor, and in each instance the females greatly outnumbered the
males. If an abundance of food tends towards the development of femaleness in

316 LIFE HISTORY OF THE AUSTRALIAN OYSTER,

this species, a preponderance of males would certainly be expected. On the
evidence, therefore, it does not appear that the food supply has an influence on the
sex-determination of O. commercidlis.

Sizes and Characteristics of Oyster Larvae at Various Stages of Growth.

During the course of the investigation on the Hawkesbury River at various
times from January to May, 1925, copious measurements were made of the oyster
larvae taken in the plankton net. The net was 12 inches in diameter and the usual
conical shape; the material used consisted of bolting silk, No. 16, containing 157
meshes to the inch, which was the finest obtainable in Sydney. Unfortunately the
mesh was not sufficiently fine to retain the early straight-hinge stages, so that
there remains a gap between the largest of those raised by means of artificial
fertilization and the smallest of those retained by the net. We have seen that the
dimensions of the largest larvae developed by artificial fertilization were: Length,
75u; depth, 584; hinge, 50u. The smallest larvae obtained in the plankton net
measured 1454 long; 1334 wide; and hinge 75u. From this size to that of the
fully grown larva a complete gradation was obtained (PI1. xxvi, fig. 34, a-c). Many
hundreds of measurements were made by means of an ocular micrometer, using a
2/3 inch objective. It was found that larvae of the same length exhibited minor
variations in depth, hinge-line or umbo, much of which may be attributed to their
lack of symmetry, which makes uniform orientation difficult. The differences,
however, were very slight, and a sufficient number of individuals was examined to
allow of a fairly accurate average being obtained.

The measurements given in the following table range, by single micrometer
units, from 18 units (1454) to 40 units (333,n).

Dimensions (in micra) of the Larvae of O. commercialis.

Total Depth
Length. including Hinge. Umbo.
Umbo.

75 58 50
145 133 75
158 142 75
166 145 83
175 159 8
182 166 16
192 175 25
200 182 25
208 192 25
216 200 25-33
226 208 33
233 216 33
242 226 33-42
250 233 42
258 242 50
266 250 58
275 258 58
283 275 58
292 283 58
300 300 58
308 312-325 58-66
312 342-350 58-66
325 350-358 58-66
333 366 58-66

~]

BY T. C. ROUGHLEY. 31

A series of measurements of the larvae of the American oyster (0. virginica)
has been given by Stafford (1918); of the Japanese oviparous oyster (0. gigas)
by Hori and Kusakabe (1926); and of the Japanese larviparous oyster
(O. denselamellosa) by Seno (1929). For purposes of comparison they are shown
in the following tables.

Dimensions of O. virginica, in micra.> Dimensions of O. gigas, in micra.
Length. Depth. Hinge. Length. Depth. Length. Depth.
69 55 48 80 68 165 185
76 62 48 90 80 180 200
83 69 48 95 94 200 220
89 76 48 100 107 220 240
96 83 48 110 120 250 265
103 89 48 120 132 270 280
110 96 48 125 140 280 290

117 96 55 143 160

124 108 55

131 110 48

138 110 55

145 ialyy 55

Wake 12¢ 55 Dimensions of O. denselamellosa, in micra.
159 131 55

165 145 55

172 148 55 Length. Depth. Length. Depth.
179 152 55

207 193 - _

241 221

276 262 148 126 325 370
310 290 235 198 347 370
345 296 280 201 372 382
379 345 295 Zoi 403 387
386 359 325 | 248

A comparison of the length-depth ratio of the larvae of the four species
shows certain interesting differences. In the case, first, of O. commercialis, we find
that, up to 300u, the larvae are longer than deep; at 300u, the length and depth
are equal; from about 300u until they set as spat, their depth is greater than their
length.

In O. virginica, it is seen that in all stages of development, from the youngest
to the fully-grown larvae, the length is greater than the depth.

A further striking contrast is seen in O. gigas, in which only the very youngest
stages, i.e., those under 100u long, are longer than their depth; from the 100u
stage till they set as spat, when they are 280u long, their depth is greater than
their length.

The measurements of O. denselamellosa, as given by Seno, show a variation so
great, compared with the very small variations in the other species, that one is
tempted to conjecture that either more than one species of Ostrea, or, perhaps, some
other bivalve larvae, have been included in the measurements; or possibly that the

1Stafford’s measurements are given in ocular micrometer units, each of which
in terms of the stage micrometer is 6:9. His units, therefore, multiplied by 6:9,
give their value in micra.

318 LIFE HISTORY OF THE AUSTRALIAN OYSTER,

orientation of the larvae has been unsatisfactory. Up to a length of 295u the
length remains constantly greater than the depth, but of the two larvae recorded
with a similar length of 325u, one has a height of 248u, and the other a height of
370u. A difference in the height of 122u in two individuals of the same species in
which the length is the same is an extraordinary variation, anything approaching
which does not appear ever to occur in O. virginica, and certainly does not in
O. commercialis. Stafford records a number of variations from his average measure-
ments, and the greatest of them is a difference in the height of 27» in a larva
whose length was 138u. The bulk of the variations, however, rarely exceed 6m. In
my own measurements, it was exceptional to find a variation greater than 8u; the
greatest variation was a difference in the height of 20u in a larva which measured
258u in length.

Summarizing the dimensions of the larvae of O. commercialis, O. virginica,
and O. gigas, it may be said that O. commercialis is longer than deep in all but the
latest stages of larval growth; O. virginica is always longer than deep; and O. gigas
is deeper than long in all but the very earliest stages of larval life.

Unfortunately, progressive measurements of O. edulis do not appear to have
been recorded; they should prove highly interesting for comparison. Boury (1930)
states, however, that they measure between 200u and 230u long when they pass
from the straight-hinge stage to the umbo stage. The proportions of a larva at the
beginning of the free-swimming stage are: length, 100; height, 90; hinge, 45.
Larvae in the straight-hinge stage varied from a length of 165u and a height of
152u, to a length of 229u and a height of 203u. In the umbo stage they varied
from a length of 2004 and a height of 183u, to a length of 275u and a height of
260u. The larvae of O. edulis, therefore, resemble those of O. virginica in the
respect that their length is always greater than their height.

We are now in a position to compare the length of fully-grown larvae of the
Australian oyster with those of other species. Marked differences obtain, as the
following tabulated list shows:

O. edulis (larviparous) .. .. .. 270u O. virginica (oviparous) SOL
O. lurida (larviparous) .. .. .. 255u O. gigas (Oviparous) .. .. .. 2804
O. denselamellosa (larviparous).. 380u O. commercialis (oviparous) ROO UL

Measurements of the thickness of larvae are not easy to obtain on account
of the difficulty of standing them on their edges. By manipulation with needles,
however, I succeeded on several occasions in leaning the larvae in an almost
vertical plane against other objects in the plankton, and obtaining measurements.
They were found to possess very considerable thickness, much greater than was
indicated when they were lying flat (Pl. xxvi, fig. 34, w). For instance, one
individual 230u long was 175 thick; another, 300% long was 208 thick; and a
third 3124 long was 216 thick.

Distribution of Laruae.

The early stages of larval oysters are captured in abundance when towing the
plankton net at the surface of the water; here, however, the catches of older larvae
decrease in proportion to their size. Fully developed larvae and those approaching
full development were found to be more abundant near the bottom.

Oyster larvae, along with other matter suspended in the water, tend to collect
in eddies, and I have repeatedly found that the best crops of oysters occur in
situations where the currents, by the contour of the land, form eddies. Striking
confirmation of this was obtained one afternoon when working on the Hawkesbury

BY T. C. ROUGHLEY. 319

River. During the morning a considerable quantity of fruit had been dumped over-
board from a lighter, and in the afternoon was found to have collected in a small
eddy opposite the boatshed in which I was working. I towed the plankton net for
10 minutes over an oyster bed some distance from the eddy, and, after washing
the contents of the net into a beaker, I returned and towed the net in the area
where the fruit was floating. The number of oyster larvae obtained in the second
haul was very much greater than that secured in the previous one.

A study of currents and of the eddies formed by bays and headlands is of very
great value in determining the location of good spatting grounds. Two of the most
prolific spatting grounds in New South Wales are situated in Salamander Bay,
Port Stephens, and in the bay east of the Hawkesbury River railway station. In
the former, the water eddies strongly on the ebb tide, and in the latter on both
flood and ebb tides.

In New South Wales the best spatting grounds are found in the estuaries near
the entrances of the rivers. Here, the salinity of the water is consistently high,
and the water contains much less sediment in suspension; consequently, material
in the water which offers a surface for the attachment of spat remains compara-
tively clean. Oyster larvae cannot attach to a muddy or slimy surface. In such
situations, however, the growth of spat is much slower than it is upstream, where
the salinity is lower on account of the fresh water continually being received from
the head of the river. As an instance of the effect of a higher salinity on the
growth of oysters, an excellent example may be cited in the Hawkesbury River.
In Cowan Creek, a tributary of the Hawkesbury River near its entrance into
Broken Bay, the oysters, although occurring in most prolific crops, remain very
stunted and rarely grow to a marketable size, even after many years. West of
the railway crossing, however, where the effect of the fresh water from the source
of the river and its numerous tributaries is continually felt on the ebb tide, growth
is very rapid. If the stunted oysters from Cowan Creek are transferred to
situations of lower salinity upstream they usually thrive and grow rapidly,
provided that they are reasonably young when removed. The old, thick-shelled
oysters from Cowan Creek rarely increase their size to any extent when transferred
to other beds.

Another striking example of the effect of a high salinity is to be found in Port
Hacking, situated a little south of Port Jackson. The amount of fresh water which
enters Port Hacking is negligible, and the oysters, although very abundant, rarely
grow to marketable size. If, however, the younger individuals are transplanted in
the George’s River, situated a few miles north, they quickly add to their shell
growth, and in the course of eighteen months or two years develop into good
marketable oysters.

An oyster larva begins to feed when it becomes a trochophore. It is then a
naked cluster of cells, and swims by means of cilia strongly developed on the
anterior surface. When the larva is first enclosed within shells the hinge-line is
straight or slightly concave. From then till the hinge is obscured by the umbos the
larva is usually referred to as being in the straight-hinge stage. At first the larva
is pale grey in colour and semi-transparent; it gradually develops a bluish tint
towards the dorsal margin. It is now provided with an alimentary canal in one
loop, consisting of a primitive mouth leading through a short oesophagus into the
stomach, which is merely a dilation of the alimentary tract, and this constricts
posteriorly to form the short intestine, ending in the anus. The digestive diverti-

320 LIFE HISTORY OF THE AUSTRALIAN OYSTER,

cula shortly become apparent as yellowish or yellowish-brown cells partially
surrounding the stomach dorsally. The ciliated end of the larva (anterior) has
now developed into a contractile velum, the cilia of which are longer and stronger
than in the embryo. The velum is withdrawn inside the shell when it is closed
and expands and projects beyond it when the shell opens to allow the larva to swim.

In O. commercialis the straight-hinge stage persists till the larva reaches a
length of about 1754. From thence onwards the hinge begins to become obscured
by the development of the umbos. In O. virginica the umbos begin to show
prominently when the larva is 1724 long (Stafford, 1913), and in O. gigas when it
is 954 long (Hori and Kusakabe, 1926). Coincidental with the development of
the umbos, another organ of locomotion, the foot, begins to show prominently
within the shell. For some time after the foot becomes visible it does not appear
to be protruded beyond the edge, but can frequently be seen to move within the
shell. It is situated behind the velum and oesophagus and anterior to the posterior
adductor muscle; ventrally it extends in an antero-posterior direction along the
line of the thickened edge of the mantle. With the development of the umbos the
opacity increases, due partly to the increased thickness of the shells and the
enclosed tissues, and partly to the deposition of pigment. The adductor muscles,
of which in the larva there are two, an anterior and a posterior, can now be
distinguished, and the digestive diverticula have developed a brownish hue.

The smallest stage at which I have seen the foot protruded beyond the shell is
when the shell is 230u long (240u in O. virginica—Stafford). From thence till
the larva settles down as spat the foot is used to enable it to crawl about.

When the larva is fully grown, and is ready to attach to some object as a spat,
its opacity has increased very greatly and it is heavily pigmented. The shell,
viewed by reflected light, is bluish-brown in colour and of a horny consistency;
it shows concentric lines of growth, the spaces between them becoming wider
towards the growing edge; they vary in number from 9 to 12. When viewed by
transmitted light, the general colour is a yellowish-brown. Closer colour analysis
shows that the dorsal region, in the vicinity of the hinge and umbos where the
digestive diverticula are situated, is very dark brown, the middle area of the larva
is yellowish-brown, sometimes with a greenish tinge, and the ventral edge pinkish-
brown. The adductor muscles are bluish-grey, and the anterior prolongation of
the foot is pink.

The mouth (Pl. xxvi, fig. 35, mo.) is situated between the velum and the foot;
it leads by way of the ciliated oesophagus (@) into the stomach (s), which lies
below the hinge and is elongated antero-posteriorly. The intestine (i) leaves the
stomach near its posterior border on the left side, immediately coils upwards and
extends anteriorly into the left umbo (l.u.), coiling downwards when near the
edge of the shell, and then extending backwards with a deep ventral loop, termin-
ating in the anus (a) situated above the posterior adductor muscle. The epithelium
of the whole of the digestive tract is ciliated, that of the stomach more strongly
than the rest. The thrust of the cilia causes the food to revolve in a counter-
clockwise direction. The dark brown cells of the digestive diverticula (d.d.) now
completely envelop the stomach, and extend dorsally into both right (r.w.) and
left (l.w.) umbos. The adductor muscles are oval in shape, the long diameter
extending vertically; the posterior adductor (p.m.) is somewhat larger than the
anterior (a.m.).

The velum (v.) occupies practically the whole of the area between the anterior
adductor muscle and the median vertical line, i.e., anterior to the oesophagus. The

BY IT. C. ROUGHLEY. BY

to

cilia are extremely long and powerful, and can be plainly seen through the shell
when it is folded at rest. When protruded and extended it opens up to form a
large, oval pad, exceeding the greatest width of the opened shells when seen
swimming towards the observer’s eye (Pl. xxvi, fig. 34, 1). The foot, when at rest,
occupies the space between the posterior adductor muscle posteriorly and the
oesophagus anteriorly, extending ventrally to the edge of the mantle (Pl. xxvi,
fig. 35, f.). It exhibits a pronounced prominence posteriorly, resembling somewhat
the heel of the human foot, and the anterior prolongation extends forward in close
proximity to the edge of the mantle, as far as the mouth. The velum is protruded
much more frequently than the foot, and usually precedes the protrusion of that
organ. Prior to the protrusion of the foot, it exhibits considerable movement
within the shell, and the tip may be frequently projected and withdrawn. It gives
the impression of great nervousness. When the whole of the foot is protruded
beyond the shell (Pl. xxvi, fig. 34, m), it stretches and becomes narrower as it
extends; the tip fastens to the bottom and a sudden contraction draws the shell
after it. By a continued repetition of this movement the larva is capable of
crawling considerable distances. When fully extended the length of the foot is
about equal to the length of the shell,

On several occasions it was noticed that the foot enabled the larva to creep
over the surface of the watch-glass in which it was being observed, by the contact
of the cilia on the base of the foot with the bottom, the movement of the larva
in this case being a slow and even progression forwards, as distinct from the
rather violent jerks which ensue from the contraction of the foot when the tip
is attached to the glass.

The foot is ciliated over the whole of the protrusible surface, the cilia along
the ventral surface being larger and stronger than those elsewhere. There is a
median groove on the ventral surface (1). The upper half of the portion pro-
truded beyond the shell is pale magenta in colour; the lower half is pale yellowish-
brown.

When the valves of the shells open the right and left mantle-folds extend
to their edges, and are withdrawn a short distance within the shell when they
close, the thickened edge of the mantle (PI. xxvi, fig. 35, mn) showing as a well
defined line extending from the anterior edge of the anterior adductor muscle to
the posterior edge of the posterior adductor.

Conspicuous in the advanced stages of larval life are two very dark, almost
black, pigment spots (otocysts, 0), one on each side about the centre of the
larva.

Toward the end of larval life the gills (g) begin to make their appearance,
and may be distinguished in the posterior half of the larva as a transversely
ridged band extending from behind the pigment spot downwards and backwards,
terminating below the posterior adductor muscle. They are broadest anteriorly,
and taper uniformly towards the posterior extremity.

When the velum and foot, or velum alone, are protruded the organs situated
above them are pulled downwards an appreciable distance, the stomach then
occupying the central region of the body.

The younger larvae are continually on the move from the bottom to the
surface of the water, but as they become older they come to the upper layers
with less frequency. Currents, however, carry them backwards and forwards and
it is impossible to say where the progeny of any oysters will settle down as
spat. Having completed its cycle as a free-swimming organism, the larva must

By Ar LIFE HISTORY OF THE AUSTRALIAN OYSTER,

come in contact with some clean, firm surface, or within a short time, stated
by T. C. Nelson (1921) to be about two days, it will usually die. Wells (1926)
has recorded that when raising oyster larvae to the setting stage by means of a
milk separator, a few were found which developed into perfectly healthy oysters
without attachment, although there was plenty of material for them to attach
to. The power of attachment appeared to be weakened or lost.

It appears to be characteristic of oysters everywhere that far greater numbers
attach to the under surfaces of objects in the water than to the upper. This
obtains in New South Wales, and care is always taken when laying out cultch
to see that it is not placed vertically. Stone slabs may be leaned one against
the other like an inverted V, or they may be stuck into the mud at an angle of
about 45°. When mangrove sticks are used they may be bunched at an angle or
laid horizontally. A reason for this peculiarity of the disposal of the young
spat may be found in the fact that the upper surfaces of submerged objects quickly
become coated with sediment, and that the heat of the sun’s rays proves too
severe for many of those spat that do attach to the upper surfaces of objects in
the tidal zone. There is abundant evidence that the heat of the sun on a hot
summer’s day does occasionally kill considerable quantities of oysters in New
South Wales, and it is only reasonable to expect that the newly attached spat,
which, up to the time when they attach themselves, have spent the whole of
their existence in the water where extremes of temperature such as are met
with on the foreshores are not encountered, will be particularly susceptible to
undue heat. Mangrove sticks, when used for the purpose of catching spat, are
always bunched closely together, and are later (usually in the spring following
attachment) transferred to beds where growth is more rapid; on these beds the
sticks are stuck into the mud in an upright position a foot or two apart. Now,
the greatest crops are always obtained on the inner layers of sticks, which are
more or less sheltered from the sun’s rays by the outermost sticks. Experience
has shown that if these sticks are separated during the hot summer months,
exposure to the sudden heat frequently proves fatal to many of the oysters, which
have had no opportunity of acclimatizing themselves to such a temperature. For
this reason, an effort is always made to transfer the sticks before the hot weather
sets in.

It is at first sight rather surprising that more larvae should attach to the
lower surface than to the upper, in view of the support which the upper surface
provides, as against the necessity of clinging firmly and securely to the lower
surface. It is not improbable, I think, that far greater numbers of fully grown
larvae do alight on the upper surface than actually attach to it, and that they
then crawl about in an endeavour to get beneath the object they alight on.
If the surface is of too great an area, they can still swim in search of a more
favourable object. In other words, I am firmly of the opinion that the larvae
to some extent at least select the location of their attachment. Frequently, when
I have laid out oyster shells to catch a set of spat, I have found that the majority
were located in depressions, such as the indentations between the teeth along
the inner margins of the shells. In such situations a certain amount of protection
against abrasion is afforded when the shell is moved by tides and currents.

Then, too, there is a considerable discrimination exercised in the choice of
material to which attachment is effected. The nature of the surface plays an
important part in regulating the attachment. For instance, if the sticks of
the Black Mangrove (Aegiceras majus), the White Honeysuckle (Banksia integri-

i)
tb
ee

BY T. C. ROUGHLEY.

folia) and the Swamp Oak (Casuarina glauca) are laid out on the same bed, by
far the greatest numbers of spat will attach to the Mangrove, which has a smooth,
firm bark, while those of the Honeysuckle and Oak are rougher and more broken.

Still another factor appears to play a part in influencing the larva to seek
the lower surface of an object for attachment. When nearing the completion of
its development it exhibits a negative phototaxis. At this stage there appears on
each side near the centre of each half of the mantle a dark pigment spot known
as the pallial eye. “This spot’, states Nelson (1925), “although unable to form
an image like a true eye, is sensitive to light. Experiment shows that larvae
which possess the pallial eye are sensitive to light, whereas those that do not
are insensitive to normal illumination. In the presence of light the ‘eyed’ larvae
of the oyster are stimulated and continue moving until they get into a shaded
place, when they become quiescent. Since the pallial eye develops within 24 hours
of the time of attachment, it follows that setting during the daylight hours is
profoundly influenced by light. Since the larvae at the setting stage are active
while exposed to light, it follows that during the day they would tend to collect
in shaded places such as the under sides of shells, and such has been found to be
the case.”

The method of fixation has been actually observed by T. C. Nelson. It had
long been maintained that the substance which cemented the larval shell to the
object of attachment was secreted by the edge of the mantle. Stafford (1913),
however, questioned the validity of this opinion, mainly because the mantle edge
was engaged in the secretion of material for shell growth, and because of the
“insurmountable difficulty in the bringing of the mantle margins far enough outside
of the shell to be applied to the proper spot”. Examination of sectioned larvae
disclosed a byssus gland, capable of secreting matter that might be used in
fixation, situated in the heel of the foot, which is capable of bringing the mouth
of the gland to the necessary area on the outside of the shell. This byssus
gland was found by Stafford to occupy, in fully grown larvae, a considerable portion
of the inside of the foot; it consists of a median and two lateral lobes, with
a main duct running through the median portion and continuing to the external
opening at the end of the heel. In the youngest spat, on the other hand, the byssus
gland is relatively inconspicuous, and its cells shrunken and collapsed. Clearly, at
the time of fixation it has functioned very actively. Stafford further reasoned that,
when secreting the cementing substance, the foot is thrust forwards and upwards
until the point of the heel, on which the duct of the byssus gland opens, comes
to the area of contact, and then the secretion is poured out, flowing between the
shell and the substratum, until fully discharged, when the foot is withdrawn.

Nelson (1924) confirmed Stafford’s contention that the cementing material
is secreted by the foot, for he was able to watch the whole process in larvae that
were crawling about on a piece of glass. He found, however, that the use of
the foot differed somewhat from Stafford’s conjecture. With the foot half extended
and pointing directly away from the shell, its distal end flattened in contact with
the glass, the left valve was held inclined to the glass at an angle of about 30
degrees. In this position the ventral edge of the shell almost touched the sub-
stratum. The ventral border of the mantle was extended until it came in contact
with the glass where it remained for two minutes, and was then withdrawn.
The foot was then slowly drawn in. “The extrusion of the mantle for a short
period”, Nelson states, “evidently aids in the quick and economical distribution
of the cementing fluid as it is poured out of the byssus gland at the ventral edge
of the left valve. The secretion hardens in less than 10 minutes.”

324 LIFE HISTORY OF THE AUSTRALIAN OYSTER,

The angle of attachment of O..commercialis corresponds with Nelson’s estimate
of the angle in O. virginica. A number of measurements with a goniometer
eyepiece showed that the median line of the spat, i.e., the line of apposition of the
right and left valves, lies at an angle varying from 27 to 32 degrees, while the
angle of the upper surface of the right valve lies at an angle of about 56 degrees
(Pl. xxvi, fig. 34, 3). The average depth of newly attached spat of O. commercialis,
i.e., the distance between the furthest edges of the right and left valves, is 216u.

A note on the distribution of the larvae of 0. edulis is given by Boury (1930),
who states that during the day the larvae are found very near the surface when
the water is calm, but that they sink to the bottom when the water is slightly
agitated. At all stages of development the distribution remains the same.

A large amount of investigation has been carried out by American workers
on the distribution of the larvae of O. virginica at various stages of their growth.
T. C. Nelson (1925), for instance, found that in Barnegat Bay, New Jersey, the
larvae rise during the flood tide and sink close to the bottom during the ebb.
During the flood tide the heavier water of relatively high density creeps along
the bottom while the lighter water of lower density remains above. In the zone
of transition from the water of higher density to that of lower density, lying on
the surface of the layer of dense water, the larvae of the oyster are found in
relatively enormous numbers. By lowering or raising the intake of the hose
used to pump the water through a plankton net a distance of a few centimetres,
the oyster larvae may increase from a few dozen to thousands per hundred litres
of water. By the beginning of the ebb tide or soon afterwards the two layers
become mixed from top to bottom, when the larvae are found in greatest abundance
close to the bottom, or actually on the bottom itself.

Prytherch (1928) found that in Milford Harbour, Connecticut, oyster larvae
are most abundant at the time of low slack water, and when the flood tide develops
a velocity of 0-6 foot per second, practically all the larvae sink to the bottom.

The majority of oyster larvae produced by the spawning beds in Milford
Harbour are found to remain and set within 300 yards of its centre. This is
accounted for by the oyster larvae remaining on the bottom during the greater
part of the larval period and limiting their swimming activities to the time of
slack water.

Setting or attachment of larvae was found to take place during low slack
water and continued until the flood tide had developed a velocity of 0:33 foot
per second, or 20 feet per minute.

Prytherch states that oyster larvae are not widely distributed by tides and
currents.

A much wider distribution by tides and currents appears to obtain with the
larvae of the Australian oyster, and their concentration does not appear to be
confined to the close proximity of the spawning beds, but rather do the larvae tend
to become widely distributed, and, as has been stated previously, to concentrate
in eddies, which may be far removed from the region of the parent oysters.
Illustrations of this may be seen in the prolific spat-catching grounds east of the
railway station in the Hawkesbury River, and in Salamander Bay, Port Stephens.
Before the introduction of mangrove stick cultivation into the Hawkesbury River
the ground referred to was bare of oysters; when, however, large quantities of
bundled sticks were laid out on the flat, a prolific crop of oysters was obtained.
In Salamander Bay, the sticks, when first laid out, were far removed from any
oyster beds, yet here again the initial crop was extremely heavy, and it has been

BY T. C. ROUGHLEY. 32

co

prolific during almost every season since, despite the fact that practically all of
the spat are transferred to other beds, several miles distant, before they have
had an opportunity of spawning in Salamander Bay.

After the attachment of the larva as a spat, shell growth becomes much more
rapid. In the early stages concentric lines of growth are clearly visible, and
the deeply convex larval shell appears to lie on the flatter spat shell like a hood.
The larval shell persists for a considerable time before becoming worn off by
erosion.

Anatomical Reorganization after Fixation.

Immediately after the larva attaches itself to become a spat, marked changes
begin to occur in its anatomy. These have been well described by Stafford (1913)
and will be only briefly referred to here. A general rotation of the soft parts
within the shell takes place rapidly. The anterior adductor muscle moves upwards
and backwards and quickly disappears; the posterior adductor moves downwards
and enlarges. Accompanying these muscle movements, the other organs of the
young spat move from left to right round the persisting adductor muscle, and
the whole axis of the spat is changed. The hinge no longer lies dorsally in
relation to the soft parts, but comes to lie anterior to them; the mouth, for
instance, instead of lying near the edge of the shell farthest removed from the
hinge, in young spat assumes a position below and behind it. As the spat
shell develops, the larval shells persist for a considerable time like a hood on
each side of the hinge. The velum and foot, no longer of use, are quickly
absorbed. Palps develop as a ventral extension of the tissues lying above the
anterior extremity of the oesophagus, and the gills extend from the base of the
palps to the postero-dorsal border. In very young spat the gills consist only of
right and left hemibranchs, the left of much greater depth than the right.

In spat which have attached to glass the movements of the animal within
the shell can be clearly followed. The pulsations of the heart are more rapid
than in adult oysters and they increase as the temperature rises. When the shells
are closed the heart ceases to beat, and the cilia of the gills cease to vibrate.
Within the mantles, right and left, the movement of the blood cells can be clearly
geen; they are not enclosed in blood sinuses, but move backwards and forwards in
a course which is fairly constant in its direction.

Occurrence of the Parasite Bucephalus in the Gonad.

During the course of my examination of the gonads of several thousand
individuals of 0. commercialis, I have found five infected with Bucephalus, the
cercaria of Gasterostomum, a trematode parasite which in the adult stage is found
in certain species of fish. Two of these oysters were taken from the Hunter River
(1927), one from the Hawkesbury River (1927) and two from Cape Hawke (1932).
I had previously, in 1924, found the parasites in the gonad of O. angasi, taken
from the Pambula River. The gonad in each case contained either no sexual
products or a few degenerating eggs.

In Europe two species of Gasterostomum have been recorded, G. gracilescens
and G. fimbriatum. According to Dollfus (1922) the former develops cercaria
(known as Bucephalus haimeanus) in the liver (digestive diverticula) and genital
organ of Ostrea, Cardium, Tapes, etc.; the cercariae leave these molluscs and
enter various species of fish, such as the haddock, whiting, cod, garfish, etc.,
where they encyst; if these fish are eaten by an angler-fish the larva emerges
from its cyst in the intestine and becomes an adult Gasterostomum. G. fimbriatum
is confined to fresh water; the larvae enter mussels (Unio, Anodonta) in which

P

326 LIFE HISTORY OF THE AUSTRALIAN OYSTER,

they develop into cercariae; known as B. polymorphus; from the mussels they enter
the rudd (Leuciscus erythrophthalmus) where they encyst, and they reach the
adult stage in Perch (Perca) and Pike (Hsox). Dollfus states that the ravages
of the worm in the European oyster are inconsiderable.

In America, J. Nelson (1914) states that Bucephalus infests about 1 per cent.
of oysters (O. virginica). It is apparently far less common in the: Australian
oyster. Just what fish form its intermediate and final hosts in Australia I am
unable to say.

Summary.

1. The position of the urino-genital clefts of Ostrea commercialis and O. edulis
is compared. !

2. The relationship of the gonad to the digestive and other organs is
described.

3. The gonoducts open into the urino-gerital cleft on each side slightly
anterior to the opening of the ureter; they are in close apposition with the
visceral nerves (right and left) and are surrounded by sphincter muscle fibres
where they enter the clefts.

4. The development of spermatogonia, spermatocytes and spermatozoa, and
of ova, from the germinal epithelium of the germinal ducts is described. As the
germ cells proliferate in the ducts, the latter evaginate into the vesicular con-
nective tissue internal to them, to form branching and anastomosing follicles.

5. As the gonad continues to develop, the ducts tend to coalesce with their
neighbours. In a mature ovary or testis the vesicular connective tissue below
the ducts is almost completely absorbed, and that lying between the ducts and
the surface epithelium becomes reduced to a very narrow layer.

6. The head of the sperm of O. commercialis is spherical and measures 2u
in diameter; the tail is about 42u long. The eggs are round, oval, or more
commonly pear-shaped; they average about 0:05 to 0:06 mm. in diameter, but may
occur up to 0:09 mm.

7. Conspicuous amongst the cells of the ciliated epithelium of the germinal
ducts are commonly found large, unciliated cells with granular contents, which
stain deeply with eosin and other acid stains. These cells have a secretory
function and may at times be seen emptying their granular contents into the
lumen of the duct. They are most numerous in spawned oysters. Their function
has not been determined.

8. The absorption of eggs and sperms which remain in the gonad after an
oyster spawns is described. Their place is taken by condensed vesicular tissue,
which then continues to grow by absorbing nourishment from the blood stream
till it assumes the large bladder-like structure of normal vesicular tissue
characteristic of this region.

9. The spawning of the oyster is described. O. commercialis was found to
spawn on the ebb tide following a high spring tide (new moon). The temperature
of the water was 68° F., and the density 1-020.

10. The rate of development of artificially fertilized eggs was found to
decrease as the density of the water was lowered.

11. At a water density of 1:005 a large proportion of the eggs remained
unfertilized, and in no case did development occur beyond the morula stage.

12. At a temperature varying from 68° F. to 70° F. and with a density of
1:021, the free-swimming stage was reached in 7 hours.

BY T. C. ROUGHLEY. 327

13. The embryos are completely enveloped in shells in 34 hours at a tempera-
ture of 68°—70° F.

14. Oyster embryos, 24 hours after fertilization of the eggs, were timed to
swim at the rate of 39 inches per hour.

15. At an average temperature of 78° F. the embryos began to swim as
trochophores in 6 hours; at an average temperature of 62° F. the same stage
was not reached until 22 hours after fertilization.

16. An attempt was made to compute the length of larval life by measuring
the larvae taken in the plankton net daily, but at no period during these
investigations was there a dominant size-group.

17. The sex-ratio of 3,000 oysters from the principal oyster-bearing grounds
of New South Wales was determined. Females were found always to predominate,
varying from 54 per cent. to 88 per cent. The average percentage was: females,
73 per cent.; males, 27 per cent. In other words, there were 2:7 times as many
females as males.

18. A sex-change is indicated in this species by the fact that practically
all, if not all, young oysters spawn for the first time as males. No opportunity
was afforded, however, of following the subsequent stability, or instability, of
the sex.

19. A total of nine oysters was found (of various ages from 1 to 3 years)
which contained both ova and sperms in the gonad. The histology of five of
these is described.

20. The determination of sex in this oyster does not appear to be governed
by the amount of food available, as has been suggested for other oviparous oysters.

21. Progressive measurements of the larvae are given, and the proportions
of the larvae at various stages of growth are compared with those of other
species whose dimensions have been recorded. Up to 300u, the larvae of this
species are longer than deep; at about 300u, the length and depth are equal; from
300u till they set as spat, their depth is greater than their length.

22. When fully grown the larva measures about 3334 (1/76 of an inch).

23. Young larvae are found in abundance at the surface of the water;
older larvae tend to remain near the bottom.

24. Oyster larvae, along with other organisms in the plankton, tend to
collect in eddies, and the best spat-catching grounds are those over which the
water is caused to eddy by the contour of the land and the tidal flow.

25. More spat attach to the under surfaces than to the upper surfaces of objects
in the water, and in water of high salinity than in water of low salinity.

26. The anatomy and movements of the larva are described.

27. The occurrence of the parasite Bucephalus, the cercaria of Gasterostomum,
a trematode parasite which in the adult stage is found in certain species of
fish, is recorded from the gonad of five individuals of this species, and of one
from O. angasi.

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, 1888.—On the Anatomy and Life History of Mollusca Peculiar to Australia.
Journ. and Proc. Roy. Soc. N.S.W., Vol. 22, 1888.

WELLS, W. F., 1925.—A New Chapter in Shellfish Culture. 15th Annual Rep. Conserv.
Comm. State of New York, U.S.A., 1925.

, 1926.—Report of Experimental Shellfish Station. 16th Annual Rep. Conserv.
Comm. State of New York, U.S.A.

YONGE, C. M., 1926.—The Digestive Diverticula in the Lamellibranchs. Trans. Roy. Soc.

Hdinburgh., Vol. liv, Part 3 (No. 15).

EXPLANATION OF PLATES X-XXVII.
Plate x.
1.—Oyster (O. commercialis) with spawn oozing from the gonoduct, situated beneath
the adductor muscle.

2.—Oyster with gonad fully developed. The branching ducts show prominently just
beneath the surface. Prominent ducts are usually indicative of femaleness, but this
eyster was a male.

3.—Oyster with fully developed gonad in which the ducts are not distinguishable at
the surface.
Plate xi.

4—An oyster after a partial spawning. The dark area of the gonad has been
drained of spawn.

5.—An oyster after a heavy spawning. The bulk of the gonad has been drained
of spawn.

Plate xii.
6.—Sections of an oyster with well developed gonad, showing its relationship to the

other organs. The gonad is seen as a whitish mass enclosing the dark digestive diver-
ticula which themselves surround for the most part the intestinal canal.

BY T. C. ROUGHLEY. 331

7.—Photomicrograph of a section of a young oyster, near the anterior extremity
of the oesophagus, in which the gonad, in the form of a series of ducts, is beginning to
develop. The ducts of the gonad surround the digestive diverticula above and on the

sides, but they do not penetrate the vesicular connective tissue between the oesophagus
and the digestive diverticula. Bouin: Whrlich’s haematoxylin and eosin. »x 40. d.d.,
digestive diverticula; d.g., ducts of the gonad lined externally with ciliated epithelium
and internally with developing germ cells; e., surface epithelium; m.c., mantle cavity;
oe., oesophagus; v.t., vesicular connective tissue.

Plate xiii.

8.—The opening of the urino-genital cleft on the right side. Bouin: Ehrlich’s
haematoxylin and eosin. x 80.

9—The opening of the urino-genital cleft on the left side. »* 80. b.m., branchial
nerve; b.s., blood space; c., canal joining mid-gut and style sac; d.g., ducts of the gonad;
h.r.g., hemibranch of right gill; i., returning loop of intestine; k.f., kidney follicles;
l.v.n., left visceral nerve; 7.v.n., right visceral nerve; m.g., mid-gut; r.m., right mantle;
s.b.c,, supra-branchial chamber; s.s., style sac; w.c., urinary chamber; w.g.c., urino-
genital cleft; v.t., vesicular connective tissue of oral process.

Plate xiv.

10.—Section through a genital duct of a developing testis. c.e., ciliated epithelium
of duct; s.c., secretory cells in ciliated epithelium; s., spermatozoa; sp.1, primary sper-
matocytes; sp.2, secondary spermatocytes; spg., spermatogonia; v.t., normal vesicular
connective tissue; v.t.°, band of condensed vesicular connective tissue immediately
external to the genital duct.

11.—Section through a genital duct of a developing testis. Bles: Ehrlich’s
haematoxylin and eosin. x 138. c.e., ciliated epithelium of duct; s., spermatozoa; s.c.,
secretory (eosinophilous) cells in ciliated epithelium; sp., spermatocytes; spg., sperma-
togonia; v.t., vesicular connective tissue.

Plate xv.

12.—Section through follicles of a developing testis. Bles: Ehrlich’s haematoxylin
and eosin. x 200. s., spermatozoa; sp., spermatocytes on border of follicle; v.t., vesicular
connective tissue.

13.—Section through a mature testis in which the genital ducts have coalesced and
the vesicular connective tissue lying between them and the surface epithelium has been
reduced to a very thin layer. The testis now consists of an enormous number of closely
packed spermatozoa, the tails of which lie in well-defined lanes. Bouin: Ehrlich’s
haematoxylin and eosin. x 90.

Plate xvi.

14.—Section of a young ovary cut vertical to the surface. Bles: Ehrlich’s haema-
toxylin and eosin. x 64. ad, genital duct lined by ciliated epithelium externally and
germinal epithelium, from which eggs are developing, internally; d.1, genital duct from
which a follicle is developing by sinking into the vesicular connective tissue; eggs are
continuing to develop from the germinal epithelium lining the follicle; e., surface
epithelium ; f., follicles lined by developing eggs; v.t., vesicular connective tissue.

15.—Section through a young ovary showing a genital duct from which three follicles
are beginning to sink into the vesicular tissue. Bles: Ehrlich’s haematoxylin and eosin.
x 120. c.e., ciliated epithelium of duct; d., cross section of genital duct in which the rapid
development of the eggs is causing them to become attenuated and to extend well into
the lumen; f., follicles developing from the genital duct and sinking into the vesicular
tissue; f., a branch of a follicle in cross section; s.c., secretory cells in the ciliated
epithelium; v.t., vesicular connective tissue.

Plate xvii.

16.—Ovary in an advanced stage of development, showing a duct in cross section.
Bles: Ehrlich’s haematoxylin and eosin. x 95. c.e., ciliated epithelium of duct; e.,
surface epithelium; f.w., wall of follicle; 0., eggs developing from the germinal epithelium
of the duct; continued proliferation has caused many of the eggs to become greatly
attenuated; a few have become detached and lie free in the lumen of the duct; o.},
eggs attached to wall of follicle.

17.—Section of a mature ovary, showing eggs attached to the walls of the follicles.
Bles: Ehrlich’s haematoxylin and eosin. x 250. f.w., walls of follicles; ., nucleus;
n.1, nucleolus; o., eggs.

332 LIFE HISTORY OF THE AUSTRALIAN OYSTER,

Plate xviii.

18.—Live unstained eggs of O. commercialis. The lighter circular area in each is
the nucleus, and in some, the nucleolus, which appears as a fine, clear spot in the
nucleus, can be distinguished. x 150.

19.—Section through genital ducts and follicles of a winter male. The ciliated
epithelium lining the largest duct externally has been almost completely replaced by
greatly distended secretory cells which project well into the lumen of the duct. Bles:
Ehrlich’s haematoxylin and eosin. x 200. s.c., greatly distended secretory cells; v.t.,
normal vesicular connective tissue; v.t.1, condensed vesicular tissue filling the duct and

the follicle which has developed from it; v.t.2, follicle filled with condensed vesicular
tissue.

Plate xix.

20.—Winter male after complete absorption of unspawned spermatozoa. The
spermatozoa in the ducts and follicles have been completely replaced by condensed
vesicular tissue. Carnoy: Ehrlich’s haematoxylin and eosin. x 60. b.s., blood space;
c.e., ciliated epithelium lining the ducts; s.c., secretory cells in the ciliated epithelium ;,
v.t., normal vesicular connective tissue; v.t.1, condensed vesicular tissue filling the lumen
of the ducts; v.t.*, follicles containing condensed vesicular tissue; v.t.3, condensed
vesicular tissue external to ducts.

21.—Winter female in which the unspawned eggs are being absorbed. Bouin:
Ehrlich’s haematoxylin and erythrosin. x 44. c¢.e., ciliated epithelium of duct; 0.3,
degenerating eggs in follicles surrounded by condensed vesicular tissue; s.c., secretory
cells in ciliated epithelium; v.t., normal vesicular tissue; v.t.2, portion of a follicle
entirely filled with condensed vesicular tissue which has replaced the absorbed eggs.

Plate xx.
22.—Section of a follicle of a winter female showing unspawned eggs in various
stages of degeneration. Bouin: Ehrlich’s haematoxylin and eosin. x 3385. o0., normal
egg, the absorption of which has not yet begun; o.', eggs partially absorbed; v.t., normal
vesicular tissue; wv.t.2, condensed vesicular tissue.

23.—Winter female after complete absorption of the eggs. The eggs in the ducts
and follicles have been completely replaced by condensed vesicular tissue. Zenker-
formol: Iron haematoxylin and eosin. x 70. e., surface epithelium; s.c., secretory cells
which have largely replaced the ciliated epithelium of the ducts; v.t., normal vesicular
tissue; v.t.2, condensed vesicular tissue; v.t.8, area of condensed vesicular tissue frequently
found scattered throughout the region lying between the genital ducts and the surface
epithelium in oysters of various stages of development.

Plate xxi.

24.—Winter female after complete absorption of the eggs. Zenker-formol: Iron
haematoxylin and acid fuchsin. x 65. b.s., blood space; c.e., ciliated epithelium of
duct; e., surface epithelium; s,c., secretory cells in ciliated epithelium; v.t., normal
vesicular tissue; v.t.1, condensed vesicular tissue filling the lumen of the duct; v.t.,
follicles containing condensed vesicular tissue. ;

25.—Genital duct and portion of a follicle of a winter female after complete absorp-
tion of the eggs. Zenker-formol: Iron haematoxylin and acid fuchsin. x 190. c.e.,
ciliated epithelium of duct; v.t., normal vesicular tissue; v.t.1, condensed vesicular tissue
completely filling the lumen of the duct; v.t.2, condensed vesicular tissue of follicle.

Plate xxii.

26.—Section of the gonad of a hermaphrodite individual of O. commercialis, contain-
ing eggs and sperms in approximately equal amounts. Spermatozoa (showing as dark
masses) are filling the lumen of the ducts, and the germinal epithelium is engaged in
egg proliferation. The follicles are filled with spermatozoa and the epithelium lining
their walls is for the most part giving rise to ova. Bles: Iron haematoxylin and eosin.
x 60. c.e., ciliated epithelium of genital ducts; d.d., digestive diverticula; e., surface
epithelium; o., eggs; s., spermatozoa; v.t., normal vesicular connective tissue; v.t.,
layer of condensed vesicular tissue external to genital ducts frequently characteristic
of this region.

27.—Portion of a section shown in Fig. 26, more highly magnified. x. 445. o., eggs;
o.1, developing eggs; n., nucleus; n.1, nucleolus; s., spermatozoa; sp., spermatocytes.

Proc, Linn. Soc. N.S.W., 1933. PLATE

Ne

Oyster (O. commercialis) with spawn oozing from the gonoduct.
Oyster with gonad fully developed. The branching and anastomosing
ducts show prominently just beneath the surface.

Oyster with fully developed gonad in which the ducts are not distinguish-
able at the surface.

xX.

Proc, Linn. Soc. N.S.W., 1933. PLAYE XI.

4. An oyster after a partial spawning.
5. An oyster after a heavy spawning.

Proc. Linn. Soc. N.S.W., 1933. PLATE XII.

6. Sections of an oyster with a well developed gonad.
7. Microscopie section showing early development of genital ducts. x 40.

XIII.

PLATE

iy a NY.
fq

ieee fz

as

80.

x

The opening of the urino-genital cleft on the right side.

8.

side.

cleft on the left

of the urino-genital

The opening

oF

Proc. Linn. Soo. N.S.W., 1933. PLATE XIV.

So2es
eo0 ;
@e

OI

®

Ge Oe
ee.
De DS

2,
&

&

~)

on?

o
2@o

os
a
(B-

10

10 and 11. Sections through genital ducts of a developing testis.

XV.

PLATE

oo
OO.

N.S.W., 19

Soc.

LINN.

OC,

»
‘

PI

eloping testis.

Section through follicles of a dev

2.

1
13.

90.

Section of a mature testis.

hak
et

Proc. Linn. Soc. N.S.W., 1933. PLATE XVI.

Aaa Y
ly

14. Section of a young ovary. x 64.
15. Follicles beginning to sink into the vesicular connective tissue. x 120.

A eee

: ee ty

Proc. Linn. Soc. N.S.W., 1933. PLATE XVII.

16. An ovarian duct in cross-section. x 95.
17. A mature ovary showing eggs attached to the walls of the follicles. x 250.

SH

Proc. Linn. Soc. N.S.W., 1933. PLATE XVIII.

18. Live, unstained eggs of O. commercialis. x 150.
19. Section through genital ducts and follicles of a winter male. x 200.

sr
i

ay
on

i
ae

oi)

Proc. Linn. Soc. N.S.W., 1933. PLATE XIX.

20. Winter male after complete absorption of unspawned spermatozoa. x 60.
21. Winter female in which the unspawned eggs are being absorbed. x 4

Proc. Linn. Soc. N.S.W., 1933. PLATE Xx.

22. A follicle of a winter female with unspawned eggs in various stages of
degeneration. x 335.
23. Winter female after complete absorption of the eggs. x 70.

Proc. Linn. Soc. N.S.W., 1933. PLATE XXI.

ho
aS

Winter female after complete absorption of the eggs. x 65.
Genital duct and portion of a follicle after complete absorption of the eggs. x 190.

DS
*hi

phys

Aes ames
(ete Be

; -. : <
Tie
Rok
1 M4 r =)° S
| «tt ehac WkIe pees
: ts Sats cS pe
pou : ‘ : Baan
Pars ey
ae : j, ed ee
J i f - ai) c=
7 : : # ite
eo )F* )
.
Ne cel 1 om ; =
i - aa
A _
- 1 j :
ay - 7

Proc. Linn. Soa. N.S.W., 1933. PLATE, KXXIT-

26. The gonad of a hermaphrodite individual of O. commercialis. x 60.
27. Portion of section shown in Fig. 26, more highly magnified. x 445

Proc. Linn. Soa. N.S.W., 1933. PLATE XXII.

Ht

28. Section of a hermaphrodite gonad. uf
29. Section of a hermaphrodite gonad. x 8

v

Proc. Linn. Soc. N.S.W., 1933. PLATE XXIV.

Larvae of several species of bivalves common in the plankton, Port

30.

Macquarie, N.S.W. x 60.
31. Oyster embryos, 20 hours old, swimming from the surface to the
bottom in vertical columns.

PEL
Piss

u 1 &
>) 5 5
7 va)
- 2
~ ie '
- ny

Proc. Linn. Soc. N.S.W., 1933. PLATE XXv.

32. Shelled oyster larvae reared from artificially fertilized eggs. x 100.
33. Larvae of O. commercialis in various stages of development. x 65.

Proc. Linn. Soc. N.S.W., 1933. PLATE XXVI.

34. Stages in the development of the oyster larva. x 72.
35. Fully-grown oyster larvae viewed (A) by reflected light, and (B) by
transmitted light. x 135.

Sages

Saye

~
ey

Proc. Linn. Soc. N.S.W., 1933. PLATE XXVII.

trim
7.7
36. A 1:5 mm. spat with shells open and mantle fully extended. x 53,

37. Section of a 1:5 mm. spat. x 40.

syste

BY T. C. ROUGHLEY. 333

Plate xxiii.

28.—Section of a hermaphrodite gonad in which the genital ducts and follicles are
filled with spermatozoa, while the germinal epithelium is giving rise to ova. In a few
situations spermatocytes are developing on the walls of both follicles and ducts. Bles:
Ehrlich’s haematoxylin and eosin. x 178. c.e., ciliated epithelium; o.1, developing ova;
s.c., secretory cells; s., spermatozoa; sp., spermatocytes; v.t., vesicular connective tissue.

29.—Section of a well developed hermaphrodite gonad in which the eggs and sperms
occur in about equal proportions. The development of an abundance of sperms has
been followed by very active egg development. In some situations, however, spermato-
cytes are continuing to develop from the walls of the follicles. Bles: Ehrlich’s haema-
toxylin and eosin. x 80.

Plate xxiv.

30.—Larvae of several species of bivalves common in the plankton at Port Macquarie,
New South Wales. Oyster larvae, which are characterized in their later stages by a
very prominent left umbo, are absent from this collection. x 60.

31.—Oyster embryos, 20 hours old, swimming from the surface to the bottom in
vertical columns.

Plate xxv.

32.—Shelled oyster larvae in the straight-hinge stage, reared from artificially
fertilized eggs. Five days old. Average measurement: length, 754; height, 584; hinge,
50u. x 100.

33.—Larvae of O. commercialis in various stages of development, from the straight-
hinge stage (top) to the fully developed larva with its prominent umbos (bottom right).
Collected in the plankton net, Hawkesbury River. x 65.

Plate xxvi.

34.—Stages in the development of the oyster larva. A, early straight-hinge stage;
B, late straight-hinge stage; C, the left umbo begins to develop; D, E, the left umbo
becomes conspicuous; F, the length and depth of the larva are now equal; up to this
stage the length was greater than the depth; G, fully grown larva ready to set; the
depth is now greater than the length; H, fully grown larva viewed from in front; I,
fully grown larva with shells apart and with ciliated velum fully extended, swimming
towards the surface; J, the attached larva (spat) showing the angle of attachment;
K, the foot partially extruded; L, the foot seen from below, showing the longitudinal
median groove; M, the foot fully extruded. x 72.

35.—Fully grown oyster larva viewed (A) by reflected light, (B) by transmitted
lighe x, 135. a., anus; a.m., anterior adductor muscle; d.d., digestive diverticula;
f, foot; g, gills; i, intestine; U.u., left umbo; mn., thickened edge of mantle; mo., mouth;
o., otocyst; oe., oesophagus; p.m., posterior adductor muscle; r., rectum; 7.w., right umbo;
s., stomach.

Plate xxvii.

36.—A 1:5 mm. spat with shells open and mantle fully distended. Owing to the
rotation of the soft parts within the shell, the front of the oyster is now uppermost. x 53.
a.m., adductor muscle; c.t., ciliated track along which material rejected by the palps
passes to the edge of the mantle; d.d., digestive diverticula; e.r.m., thickened edge of
right mantle; h., heart; i., intestine; l.g.l., lamella of left gill; l.l.s., left valve of larval
shell; L.p., labial palps; m.m., muscles to mantle edge; oe., oesophagus; 7., rectum;
r.g.l., lamella of right gill; r.l.s., right valve of larval shell; sh., shell of spat; s.s., style
sac; st., stomach; t.l.m., tentacles of left mantle; t.r.m., tentacles of right mantle;
v.g., visceral ganglion.

37.—Horizontal section of a 1-5 mm. spat. The front of the spat is at the right of
the photomicrograph. Bouin: Ehrlich’s haematoxylin and eosin. x 40. a.m., adductor
muscle; d.d., digestive diverticula; g., gills; h., heart; 4, intestine; l.p., labial palp;
m.g., origin of mid-gut from stomach; mn., mantle; oe., oesophagus; r., rectum; u.c.,
urinary chamber; s., stomach; s.s., style sac; v.g., visceral ganglion.

THE GEOLOGY OF THE SOUTH COAST OF NEW SOUTH WALES, WITH
SPECIAL REFERENCE TO THE ORIGIN AND RELATIONSHIPS
OF THE IGNEOUS ROCKS.

By Ipa A. Brown, D.Sc., Linnean Macleay Fellow of the Society in Geology.
(Plate xxviii; four Text-figures.)

[Read 27th September, 1933.]

1. Introduction.

2. Sedimentary Record: (a) Cambrian (?); (bv) Ordovician; (c) Silurian; (d)
Devonian; (e€) Kamfilaroi; (f) Tertiary and Post-Tertiary. .

3. Igneous Rocks: (a) Pre-Devonian; (b) Devonian; (c) Kamilaroi; (d) Tertiary.

4. Tectonic History and Igneous Activity.

5. Magmatic Differentiation.

6. Summary and Conclusions.

7. Bibliography.

1. Introduction.

The study of the geology of the South Coast of New South Wales reveals an
interesting variety of sedimentary formations, ranging in age from lower
Palaeozoic to upper Cainozoic, with which are associated igneous rocks of sub-
alkaline, monzonitic and alkaline facies. Although the geological record is far
from complete in this area, it is sufficiently well-preserved to show a definite
relationship between the history of tectonic movement and igneous injection, and
also the evolution of a series of igneous rocks, which in general gives evidence of
progressive magmatic differentiation throughout the ages, commencing from
Devonian or even earlier times.

The area is not a complete unit in itself; nevertheless it may be regarded as
a fairly typical section across the margin of a continental mass, which has been
growing by repeated deposition of sediments along its borders and their subsequent
compression and elevation to form portion of the land-mass. Throughout the ages
from early Palaeozoic to late Mesozoic time there has been a gradual change in the
position of the axis of geosynclinal deposition and in the corresponding directions
of folding and faulting of the adjacent sedimentary formations.

It is proposed therefore to give in the following pages a brief outline of the
geological history of sedimentation and tectonic structure in the area under
consideration and to attempt to show their possible relationships to the processes
of magmatic differentiation and injection of the associated igneous rocks.

The paper is a revision of portion of a thesis accepted by the University of
Sydney for the degree of Doctor of Science, and is based largely on the writer’s
previous studies of the igneous and sedimentary rocks of the South Coast,
descriptions of which have been published in These Procereprnes and in the Journal
of the Royal Society of New South Wales.

ce
co
ol

BY IDA A. BROWN.

2. SEDIMENTARY Recorp.
(a). Cambrian(?).

The oldest bedded rocks of the region outcrop from the head of the Clyde River
to the Victorian Border, a distance of 150 miles, and from the coast inland for a
distance of about twenty miles. To the west they are bounded by granitic
intrusions and by Upper Ordovician and newer sediments. No detailed account
of this series has yet been recorded, but it is hoped that such may be given in a
later paper.

These rocks consist of a series of black, banded radiolarian cherts, and meta-
morphosed fine-grained sediments, now converted into phyllites and schists. The
cherts are turquoise-bearing, but no fossils other than radiolaria have been
discovered as yet. The series is highly folded, the axes of the folds being in a
meridional direction in the northern and southern portions of the area, while in
the central portion, in the vicinity of Narooma, the fold axes are in the direction
north-north-west and south-south-east, apparently pitching to the south-south-east.
Intense minor folding and crumpling superimposed on the larger folds may be seen
in a number of cliff-sections between Bateman’s Bay and Bermagui.

No palaeontological evidence of the age of the series is yet known; the
field-relationships of this series and fossiliferous Upper Ordovician east of Quaama
indicate unconformability and suggest a pre-Upper Ordovician age for the unfos-
siliferous series. The extraordinary lithological characters of the series definitely
distinguish it from the Upper Ordovician and newer formations occurring else-
where in New South Wales. On the other hand these characters are remarkably
similar to those of certain types of Cambrian rocks in Victoria, such as those of
Heathcote (EH. W. Skeats, 1908) and the phosphate-bearing rocks of the Mansfield
district (A. M. Howitt, 1923). There are also resemblances to portions of the
Brisbane ‘‘schist” series of Queensland, particularly the Neranleigh and Bunya
Series (A. K. Denmead, 1928), and also to the pre-Silurian rocks of Cobar,
Canbelego and possibly the Forbes-Parkes districts (HE. C. Andrews, 1913a, 1913),
1910).

In the absence of more definite information, the series may be regarded
tentatively as of middle or upper Cambrian age.

(b). Upper Ordovician.

No Lower Ordovician rocks are known to occur on the South Coast, although
Upper Ordovician beds outcrop two miles east of Quaama, along the eastern side
of Pipeclay Creek, a tributary of the Murrah or Dry River. They occur immediately
east of the granite contact, and consist of quartzites and slates from which Upper
Ordovician graptolites have been obtained. These have been identified by Mr. W. S.
Dun as Diplograptus and Climacograptus. Similar forms occur in weathered slate
eight miles east of Cobargo, on the road to Bermagui, again not far from the
granite contact.

About three-quarters of a mile north-west of Narira Trigonometrical Station,
north of Cobargo, similar black slates and quartzites outcrop, and from these
Mr. C. F. Laseron collected and identified Diplograptus foliaceus, Climacograptus,
Dicellograptus (?) gracilis, D. affinis (W. R. Browne, 1914).

In each of these localities the beds are steeply dipping, and strike in a north-
westerly direction. These are the only outcrops known to occur on the South Coast
whose age can be assigned definitely to the Upper Ordovician, the main develop-
ment of these formations being in a more or less meridional zone to the west of

336 GEOLOGY OF THE SOUTH COAST OF NEW SOUTH WALES,

the area under consideration, extending from eastern Victoria across the border
to the counties of Auckland and Wellesley, New South Wales (J. E. Carne, 1897),
through the Cooma district (W. R. Browne, 1914), and northwards to the middle
Shoalhaven district (W. G. Woolnough, 1909).

(c). Upper Silurian.

The chief development of Silurian rocks is also to the west, the only known,
outcrops of Silurian being in the north-western portion of the area under con-
sideration, in the Bendithera-Wyanbene district, near the head of the Deua River.

Here fossiliferous limestones are associated with quartzites, slates and volcanic
rocks. The limestones are marmorized to such an extent that the structures of the
‘enclosed fossils are partly destroyed; the fossils identified up to the present suggest
a Silurian age, although it is possible that further collecting may prove them to be
Middle Devonian.

At Bendithera the main trend of the beds is in a north-westerly direction, and
they are faulted against, or unconformably overlain by, sediments of probable
Upper Devonian age, as described previously by the writer (1930a).

(d). Upper Devonian.

The zone of sedimentation during the Lower and Middle Devonian followed
approximately that of the preceding Silurian period, and apparently did not extend
as far east as the South Coast of New South Wales. Thus the Lower Devonian is
represented by the Snowy River Porphyries in Victoria, and by the igneous series
at the base of the Devonian in the Yass District in New South Wales; the Middle
Devonian includes the limestone series of Buchan, Bindi and Limestone Creek, and
the shales and limestones of Tabberabbera in Victoria (EH. W. SkKeats, 1929; A. W.
Howitt, 1874-1877), the Yass series of limestones and shales (L. F. Harper, 1909),
and the limestones and shales of Tarago and Lake Bathurst Railway Station in
New South Wales.

The succeeding Upper Devonian epoch is well represented on the South Coast
and has been described by the writer in a paper on the Devonian and older
Palaeozoic rocks of the area between the Shoalhaven River and the Victorian
border (1930a), and in a later paper (1931) on “The Stratigraphical and Structural
Geology of the Devonian Rocks of the South Coast of New South Wales’, in which
the relations to other Devonian formations in south-eastern Australia are
considered.

The Upper Devonian formations of the South Coast occur as a series of dis-
continuous outcrops extending from the coast in the vicinity of Twofold Bay in a
north-westerly direction through Nerrigundah, west of Moruya to the Clyde
Mountain and thence in a north-north-easterly direction towards Yalwal and the
middle Shoalhaven River.

The formation always overlies the pre-Devonian sediments with a marked
unconformity, and it has been divided by the writer (1931) into three stages,
(i) the lower or Hden stage, consisting entirely of flows of acid volcanic rocks,
400 to 500 feet in thickness in the Eden district, and somewhat thicker (800
feet) a few miles west of Moruya; (ii) the middle or Yalwal stage, including red,
freshwater shales and sandstones with interbedded volcanic flows of rhyolites and
amygdaloidal basalts allied to spilites; and (iii) the Upper or Lambie stage, about
1000 feet in thickness, of shallow-water marine conglomerates, sandstones,
quartzites and shales.

BY IDA A. BROWN. 337

In the Eden district the Upper Devonian beds have suffered relatively little
folding since their formation. The intensity of the folding increases somewhat
towards the north, but is never great except in the vicinity of igneous intrusions,
as at Araluen and Yalwal (E. C. Andrews, 1901). The axes of structural folding
are parallel to the main trend of the outcrops.

(e). Kamilaroi.

Outcrops of Kamilaroi sediments are confined to the area north of Bateman’s
Bay, and have been described in detail by L. F. Harper (1915) in his Memoir on
the Southern Coalfield of New South Wales. The Lower Marine Series and the
underlying Carboniferous, which outcrop in the Hunter River district, are not
exposed in the Southern Coalfield. Thin seams of coal belonging to the Greta
Series have been identified near the head of the Clyde River and in tributaries
of Wandandian Creek. ;

The Upper Marine Series attains a thickness of over 2,000 feet on the South
Coast. The basal beds are the Ulladulla Mudstones, which have a somewhat local
distribution. They are richly fossiliferous and contain evidence of contem-
poraneous glacial conditions in the form of ice-carried erratics and ‘“glendonite”
pseudomorphs (T. W. E. David and others, 1905; E. O. Thiele, 1903; L. F. Harper,
1915; I. A. Brown, 1925a). The Nowra grits contain a basal phase of Conjola
conglomerate, which rests unconformably on the older Palaeozoic sediments and
granite of probable late Devonian age in the neighbourhood of Conjola. The
Nowra grits are overlain by crinoidal shales, which outcrop on Nowra Hill. The
close of the Upper Marine Stage is represented by the extensive development of
red tuffs and interbedded lava flows of the Illawarra District (L. F. Harper, 1915),
and the hypabyssal intrusions of the Milton district (L. F. Harper, 1915; I. A.
Brown, 1925D).

The overlying Upper Coal Measures outcrop along the eastern slopes of the
Illawarra Tableland north of the Shoalhaven River and form the sea-cliffs between
Thirroul and Stanwell Park. They dip slightly to the north and west, and underlie
the Triassic sediments of the Sydney District.

(f). Tertiary and Post-Tertiary.

Tertiary formations occur at intervals along the coast south of the Shoalhaven
River and consist chiefly of horizontally-bedded sandy sediments of lacustrine or
estuarine origin, in which fragments of fossil wood frequently occur (C. Barnard,
1927). On the coast south of Corunna Lake, ferruginous grits unconformably
overlie Lower Palaeozoic cherts, and from these Upper Cainozoic shells have been
identified by Mr. F. A. Singleton, M.Sc., of Melbourne University. These are the
only known marine Tertiary of eastern New South Wales.

Partially consolidated sandstones, grits, conglomerates and peaty beds,
altogether nearly 100 feet in thickness, form old sea-cliffs behind the beaches
between Bournda Island and Twofold Bay. In numerous localities there are
deposits of old river gravels 40 or 50 feet above the present river level, and along
the coast are old beach deposits now above the reach of high water or storm waves.
The correlation and determination of the ages of these deposits is somewhat
difficult and much work remains to be carried out on this problem.

Sometimes olivine-basalts are associated with the Tertiary sediments as
volcanic flows, sills and dykes. Those occurring close to the coast probably were
never of great thickness, but petrologically they are similar to the basalts of
the Southern Tablelands which occur at Mt. Darragh, Tantawanglo, Brown
Mountain and Robertson.

338 GEOLOGY OF THE SOUTH COAST OF NEW SOUTH WALES,

The contact metamorphism of Tertiary sandstones by the coastal Pliocene
basalts has produced flinty quartzites that are quarried as “silica” for the manu-
facture of refractory bricks (I. A. Brown, 1925c).

3. THE IGNEOUS ROCKS.

The igneous rocks of the South Coast of New South Wales and the adjacent
tableland form an association which shows a progressive variation in chemical
composition throughout geological time. It is considered, therefore, that a study
of their occurrence may throw some light on the processes of magmatic differen-
tiation, the relationship between the various branches of igneous rocks, subalkaline,
monzonitiec and alkaline, and the relation between igneous injection and tectonic
structure.

Although the determination of the exact geological ages of many of the igneous
occurrences is a difficult matter, it is almost certain that the chief extrusions and
intrusions of subalkaline character are Devonian or older, the monzonitic rocks
are probably Permian, and the relatively alkaline series of hypabyssal and volcanic
rocks may be referred to the Tertiary period.

The following descriptions summarize the principal features of the field-
occurrence and the petrology of the igneous rocks of the South Coast and adjacent
districts.

(a). Pre-Devonian.

Voleanie igneous rocks occur as flows and tuffs among the Lower Palaeozoic
sediments of Bateman’s Bay, Moruya and Narooma. This series is tentatively
regarded as equivalent to the Cambrian Heathcotian Series of Victoria. No detailed
work has yet been carried out on these South Coast rocks, so that their affinities
are not known with certainty.

A few dykes and thin flows of volcanic igneous rocks occur among the Upper
Silurian shales and limestones of the Bendithera district, west of Moruya, but
these have not been subjected to special investigation.

(b). Devonian.
(i). Voleanie Series.

During the Devonian period in south-eastern Australia extensive flows of acid
lavas occurred over the pre-Devonian surface, along zones which were being sub-
jected to downwarping prior to the deposition of lacustrine and marine sediments.
These extravasations occurred during the Lower Devonian (?) in the Snowy River
District of Victoria, and continued into the early Upper Devonian along a zone
through Gippsland, Victoria, the south coast of New South Wales, and the Mudgee
district. During the early Upper Devonian the rhyolites were accompanied by
flows of amygdaloidal basalt or melaphyre. It will be shown later that probably
the acid and basic flows were both consanguineous with the granodiorites that were
intruded subsequently as batholiths along the South Coast and Southern Tablelands.

The petrological and chemical characters of the Devonian volcanic rocks of the
South Coast have been described by the writer (1931), and the Victorian rocks with
which they have been correlated are described by A. W. Howitt (1874-76), BE. W.
Skeats (1909, 1929), H. O. Teale (1920) and others.

The Upper Devonian acid lavas of the South Coast are normal potassic
rhyolites, which show slight lithological variations due to differences in the
conditions of consolidation. The stratigraphically lower members of the series are
usually porphyritic in quartz, and have a stony groundmass, showing devitrification

BY IDA A. BROWN. 339

in thin section. In spite of the relatively high percentage of potash, the felspar
crystals, when present, often consist of acid plagioclase, and it is assumed that the
potash is present in the groundmass. The upper horizons of rhyolite are character-
ized by well-marked banding and spherulitie structures.

Table 1 illustrates the chemical composition of the South Coast rhyolites and
their equivalents in the adjacent area of Victoria. Further reference will be made
to them when considering the magmatic differentiation of the igneous rocks.

TABLE 1.
ie Ine IIl. IV. Vi VI. VII.

SiO, 75°34 75-91 78°64 78°47 72°55 62°56 74-72
Al,O; 11-89 11:89 9-85 10-68 11-74 16-60 13°05
Fe.0; 1-54 1-58 0-54 0-18 2-54 1-02 0-52
FeO 1-60 0-96 2-00 2°23 0-46 5-98 1-42
MgO 0-28 0-47 0-10 tr. 0-68 Ace (AL 0-41
CaO 0-16 0-26 0-80 0:66 1-85 4:3 0-66
Na,O 2:06 22.3 2-03 3:29 3°46 2-98 3°62
K,0 3°82 5:59 5-16 4-15 4°41 PAO 4°31
H,O + 1-18 0-58 0-40 0:2 0-41 0°68 0-61
H,O — 0-16 0-09 0-14 0-09 0-06 0-18 0-13
CO, 1:60 = = = 1-80 — 0-08
TiO, 0:31 0-28 0:67 0:59 0-175 1-10 0-16
P.O; = = — tr tr 0-14 0:17 0-38
MnO tr. tr —_— — _ tr —

Total ae ae 100-04 99-84 100-33 100-54 100-27 100-85 100-07

I. Rhyolite, Quarry, east of ‘“Edrom’’, East Boyd, Twofold Bay. Anal. I. A. Brown,
Proc. LINN. Soc. N.S.W., lIvi, 1931, p. 476.

II. Rhyolite, Deua River, road to Araluen, 11 miles from Moruya. Anal. I. A. Brown,
Proc. Linn. Soc. N.S.W., lvi, 1931, p. 476.

Ill. Banded Rhyolite, southern plateau of Wellington, Victoria. Anal. E. O. Thiele,
Proc. Roy. Soc. Vict., xxi, 1908, p. 266.

IV. Quartz-porphyry, southern shore of Lake Karng, Wellington, Victoria. Anal.
G. Ampt, Proc. Roy. Soc. Vict., xxi, 1908, p. 266.

V. Quartz-porphyrite, No. 100, Mt. Tara Ranges, Snowy River Porphyry Series.
Anal. HE. O. Teale, Proc. Roy. Soc. Vict., xxxii, 1920, p. 125.

VI. Dacite, Willimigongong Creek, near “‘Cheniston’’, Upper Macedon, Vict. Anal.
Lewis and Hall, Bull. Geol. Surv. Vict., No. 24, 1912, p. 17.

VII. Rhyolite, Blue Hills, Taggerty, Vict. Anal. E. S. Hills, Proc. Roy. Soc. Vict.,
xli, 1929, p. 189.

The associated basalts usually have an altered appearance and for this reason
are commonly called ‘“‘melaphyres’’ in Victoria. The writer has shown (1931,
p. 478) that they have some affinities to the spilites and that the felspar is usually
an acid plagioclase. Although many specimens contain unaltered augite, the rocks
contain an abundance of alteration products which may be regarded as deuteric.
These include calcite, epidote, chlorite and zeolites, which occur in the groundmass
and in the amygdules, and the rock may be traversed by veins of fibrous asbestos
and quartz. The following analyses are typical of the basic flows in the Devonian

rocks of south-eastern Australia.

340 GEOLOGY OF THE SOUTH COAST OF NEW SOUTH WALES.

TABLE 2.

I II Tit
SiO, 46-28 49-87 49-35
Al,O; 16-02 15-91 17-61
He;O3.- 2-43 3°55 1:50
FeO .. STEMS 10-09 9-72
MgO 6:84 4-84 Bor?
CaO .. 8-86 8-27 aera
Na,O 2-83 PAP AY/ 3°10
GO) a6 0-25 1-10 1-56
H,O+ 3°65 2-44 2-56
H,O— 0:30 0-19 0:65
Co, 2-58 0-22 —_
TO. 1-94 1-89 2-83
P.O; 0-34 —_ tr
MnO ae sie 0-11 pr. 0:07
Other Const. ae 0-34

Total .. o6 99-70 100-54 100-17

I. Amygdaloidal Basalt, east of Nethercote, Eden District, N.S.W. Anal. I. A. Brown,
Proc. LINN. Soc. N.S.W., lvi, 1931, p. 479.

II. Compact Basalt, Por. 68, Par. Eden, N.S.W. Anal. I. A. Brown, Proc. LINN.
Soc. N.S: W.,, lvi; 1931, p. 479.

III. Melaphyre, Moroka Snow Plain, Victoria. Anal. G. Ampt, Proc. Roy. Soc. Vict.,
p:@:0.40 i 7-1 | a ORES oe

(ii). Plutonic Series.

After the deposition of the Upper Devonian Series of sediments, extensive
batholithic intrusions took place in central and south-eastern New South Wales,
brief accounts of which have been given by W. R. Browne (1929) and H. W. Skeats
(1930). So far no detailed petrological investigations of the large batholiths have
been recorded.

Within the region under consideration there occurs a large granodioritic batho-
lith, some 25 miles in width and at least 100 miles in length, which extends from
eastern Victoria in a northerly direction through Towamba, Wyndham, Bemboka
and Bega to beyond Cobargo. It does not outcrop on the coast, but extends from a
few miles inland near Bega in a westerly direction towards Bombala and
Nimmitabel on the Monaro Tableland. For convenience this intrusion may be
called the Bega batholith, after the largest town on its outcrop.

Smaller granodioritic intrusions occur at Moruya, Coondella, Merricumbene
and Nelligen; the Braidwood-Araluen intrusion is possibly connected with the Bega
batholith by way of the Monaro tableland.

The Bega Batholith.

The boundaries of the Bega Batholith are indicated on the accompanying
sketch map (Plate xxviii).

The intrusion is composite and several distinct but probably comagmatic
rock-types may be distinguished: it is therefore possible that the injection of the
batholith may not have taken place in a single act, or even during a single
orogenic epoch, but little evidence is available on this problem, as contact with the

BY IDA A. BROWN. 341
sedimentary rocks is usually obscure. The intrusion as a whole cuts indiscrimin-
ately across the Lower Palaeozoic series; the granodiorite east of Quaama has
intruded and metamorphosed fossiliferous Upper Ordovician and also Upper
Devonian sediments. Near Wolumla, J. BE. Carne (1897, p. 162) found evidence
of the intrusion of the Wolumla granite into gently folded Upper Devonian sedi-
ments, and similar relations occur between granite and Upper Devonian beds at
Upper Brogo and Bega.

Upper Devonian sediments overlie the granite north of Numbugga, a dairying
settlement west of Bega, but the age relationships of the formations have not been
examined. A granite at Bunnair Creek, Conjola, petrologically similar to one phase
of the Bega batholith, is overlain by Upper Marine (Permian) sediments. Tertiary
basalts occur over the eroded surface of the granodiorite on the eastern margin of
the Monaro Tableland, on the slopes of the Brown Mountain, Tantawanglo
Mountain and Mt. Darragh.

The age of at least portion of the intrusion is therefore between Late Devonian
and Permian, and quite probably it belongs to the orogenic epoch at the close of the
Devonian, known as the Kanimbla Epoch in New South Wales.

Although the batholith is essentially granodioritic in composition, more acid
and more basic plutonic types are also developed, as well as numerous pegmatitic
and aplitic veins and dykes, and a variety of basic dykes. Some of the more acid
phases are associated with ore deposits of molybdenum, bismuth, tellurium,
selenium, gold, silver and iron (E. C. Andrews, 1916, p. 157), and in this respect
resemble the younger intrusives of the New England district (E. C. Andrews,
1905-09).

The most basic phases are the gabbros and diorites of Brogo and Quaama,
which occur as inclusions of varying dimensions in more acid phases of the
intrusion, and which show signs of contact-metamorphism as a result. The freshest
specimens are diorites consisting essentially of basic plagioclase and hornblende.

The most widely distributed rock-type is one which is mineralogically compar-
able to the Moruya granodiorite; this type is common in the Cobargo district, and
along the eastern portion of the batholith through Quaama and Bega. A more
coarsely crystalline variety of the same rock forms the western exposure of the
batholith along the lower slopes of the Monaro Tableland, outcrops occurring at
Mt. Darragh, Tantawanglo, the Brown Mountain, and westwards towards
Nimmitabel. The rock is a hornblende-biotite-granodiorite or tonalite consisting of
quartz, plagioclase (Ab,;An,, to Ab,,An,,), orthoclase, with microperthite, horn-
blende, biotite, sphene and minor accessory minerals.

The rock grades into a more acid phase by a decrease in the amount of
hornblenue and the presence of more acid plagioclase. This phase is a biotite-
granite, whic. is usually porphyritic in felspar. The phenocrysts may be as much
as two and a half inches in diameter and sometimes are so closely packed that the
weathered rock resembles a conglomerate. In plan the outcrop of the porphyritic
biotite-granite occurs within that of the granodiorite; it extends from near Yourie
southwards through the Murrabrine Mountain to the west of Quaama and Bega.
Good outcrops occur between Bega and Bemboka and surrounding Candelo,
Wyndham, Towamba and Pericoe. The rock contains phenocrysts of orthoclase
with microperthite, and the coarsely crystalline groundmass consists of quartz,
orthoclase, anorthoclase and acid plagioclase of the composition Ab-,,An,., with
biotite and minor accessory minerals. Some varieties of the rock contain

342 GEOLOGY OF THE SOUTH COAST OF NEW SOUTH WALES,

muscovite, the rock thus grading into a two-mica granite in the vicinity of
Towamba and Candelo.

Border phases of the batholith at Dr. George Mountain and near Yourie are
aplitic granites containing little or no terromagnesian minerals, and muscovite-
granite occurs at the Whipstick or Jingera Mines east of Wyndham. This
muscovite-granite consists of quartz, orthoclase, soda-microcline or anorthoclase,
microperthite, albite and muscovite. Masses of red garnet occur spasmodically in
the granite, which contains pipes carrying ores of bismuth and molybdenum.
There is a similar occurrence on the western side of this batholith, to the south of
Tantawanglo Mountain (EH. C. Andrews, 1916, pp. 150-166, 143-146).

No investigation of the chemical composition of the batholith as a whole has
yet been carried out, but Table 3 contains analyses of certain of the principal
rock-types. The first three are analyses of acid granites from Whipstick, and are
interesting in that they show that a large proportion of the orthoclase and anortho-
clase must be of a sodic character. The Gabo Island granite (column iv) is
probably comagmatic with the granites of the Bega batholith. The Brogo granite,
whose analysis is quoted in column v, is a type of porphyritic biotite-granite
containing scattered large phenocrysts of orthoclase. Granites from Braidwood,
Nimmitabel and Kiandra are quoted for comparison as they are probably all of the
same age.

TABLE 3.
t
I. JOG IIl. mve Vv. VI. VII. VIII. IX. Xx.

SiO, 87-66 83°54 75-02 72-49 69-44 69-38 66°58 65°94 65-72 45-15
Al,O3 7-32 9-13 UBIOU lL 13-48 12-10 12-56 14°36 15-10 17-63 13-40
FeO; 0-30 0-50 1-40 1:16 4-65 2-40 ab8} 1-20 0-42 2-97
FeO abs 0-36 0:27 2-09 2-13 Zio 3°19 2:34 2-80 13-19
MgO 0-08 0-30 0-04 0-49 M152 1:47 1-70 Diels 1157/83 4-23
CaO abs 0-46 0-12 iLoBisl 3-03 Boy) 4°18 4°48 4-36 9-79
Na,O 3°58 3°51 3:29 3:38 2°57 OUT 3:09 3:09 3:14 2°47
K.0O 0-20 1:14 4°78 4-06 3:00 3°75 B5837/ 8397/33 2-12 0-49
H,.O+ 0-02 1:14 1:17 0-76 0-65 1:05 0-79 0-80 1-03 0:39
H,O — 0-36 0-10 0:07 0-18 0-19 0:09 0-17 0-11 0-06 2-41
CO, 0-05 abs abs. tr. = 0-01 0-04 0-12 abs. —
TiO, abs tr. 0-30 0-46 0:55 0-52 0-65 | 0°55 0-41 5-04
ZrO. Bs abs abs. abs. — abs abs. abs 0-19 =
P.O; aes 0-02 0-05 0-03 tr. 0-32 0-12 0-10 0-21 0:19 0-12
S(FeS,) .. 0:16 abs. abs. — abs. — abs. 0-13 —
MnO ne tr. abs. 0-02 0:13 — 0:04 0:07 0:04 0:08 0-19
BaO we tr. abs. abs. = — 0-10 0-04 0:13 abs —
Etc. Se 0-54 0-08 — — — = — — = =

Total .. | 100-29 | 100-31 | 100-22 99-99 | 100-15 | 100-23 99-86 | 100-37 | 100-01 99-84

Sp. Gr. 2-646 2-666 2°633 2-635 — 2-710 2-718 2-725 2-729 —

I. Granite, Whipstick, N.S.W. Anal. H. P. White, Dept. Mines, Geol. Surv. N.S.W.,
Min. Res. No. 24, 1916, p. 150.

BY IDA A. BROWN. 343

II. Biotite-Granite, Whipstick, N.S.W. Anal. W. A. Greig, Dept. Mines, Geol. Surv.
N.S.W., Min. ‘Res. No. 24, 1916, p. 150.

III. Muscovite-Granite, Whipstick, N.S.W. Anal. H. P. White, Dept. Mines, Geol.
Surv. N.S.W., Min. Res. No. 24, 1916, p. 150.

IV. Granite, Gabo Island. Anal. J. Watson, Proc. Roy. Soc. Vict., xxvi, 1914,
p. 268.

V. Granite, Porphyritic, Brogo. Anal. A. A. Pain, in Washington’s Tables, 1917,
p. 102.

VI. Granite, 6 miles north-east of Braidwood, N.S.W. Ann. Rept. Mines Dept.

NES W., 1909, p: 198.

VII. Biotite-Granite, Kybean Road, north of Bega Road, east of Nimmitabel.
Proc. Linn. Soc. N.S.W., xxxiv, 1909, p. 314. (Also Ann. Rept. Mines Dept. N.S.W.,
1908, p. 184.)

VIII. Biotite-Granite, 7 miles south-east of Kiandra (not Kiama), N.S.W., Ayn.
Rept. Dept. Mines N.S.W., 1909, p. 198.

IX. Granodiorite, 2 miles east of Moruya, N.S.W. Anal. I. A. Brown, Proc. LInn.
Soc. N.S.W., liii, 1928, p. 162.

xX. Gabbro, Brogo, N.S.W. Anal. A. A. Pain, in Washington’s Tables, 1917, p. 662.

The only small granitic intrusion whose petrology has been studied in detail
is that at Moruya (I. A. Brown, 1928), but field-observations and brief examination
of hand-specimens and thin sections of the rock-types of the other intrusions
indicate that the Moruya series is representative of the smaller granitic intrusions.
The Moruya plutonic series, with its associated dyke-rocks, forms a complete and
typical subalkaline igneous complex. The principal plutonic types are _ biotite-
granite, granodiorite, tonalite and diorite-gabbro, whose occurrence and relation-
ships have been described in detail by the writer (1928). Aplitic and lamprophyric
dyke-rocks are closely associated with the intrusion.

The other small intrusions appear to consist essentially of types comparable
to the more acid phases of the Moruya complex. This feature may be due to the
fact that they have not been sufficiently eroded to expose the more basic types
which, even in the case of the Moruya intrusion, form only a small part of the
whole. Chemical analyses of the chief types are quoted in Table 4.

(c). Kamilaroi.

Rocks of a monzonitic character occur in the northern part of the area under
consideration, where three main groups have been studied in detail. These
comprise (i) a series of volcanic flows of latite occurring chiefly in the Illawarra
District, north of the Shoalhaven River, (ii) hypabyssal intrusions of monzonite-
porphyry in the Milton District, and (iii) a monzonitic igneous complex in the
Mount Dromedary district.

(i). In the Illawarra District, volcanic flows of monzonitic composition are
interbedded with tuffs in the Upper Marine Series of Permian age. Their
occurrence has been mapped and described by J. B. Jaquet (1905) and L. F.
Harper (1915), and petrological descriptions of the rocks have been given by
G. W. Card (1915) together with chemical rock-analyses by officers of the Depart-
ment of Mines, New South Wales. Seven distinct flows have been recognized; they
all occur within 15 miles of the present coast-line “and conform to the dip of the
sedimentary rocks in every case” (Harper, 1915, p. 290). Although the rocks show
marked mineralogical and textural variations, the relative uniformity of the
chemical composition indicates that the flows are all consanguineous. They are
described by Mr. Card as latites and latitic dolerites, and are usually black rocks of
basaltic appearance, containing phenocrysts of plagioclase (generally labradorite)
in an aphanitic groundmass. Under the microscope the groundmass is seen to be
holocrystalline, and to consist of augite, plagioclase, orthoclase and minor

344 GEOLOGY OF THE SOUTH COAST OF NEW SOUTH WALES,

TABLE 4.
Ip Tike Tut, IV. Vi
SiO, a ae Bs 49-38 50-44 61-44 65°72 70-78
Al,O3 13°89 17-05 17:61 17-63 15:77
Fe,0, 1:89 2-43 1-86 0-42 0-69
FeO 6-03 5:24 3-59 2-80 2-44
MgO 15-82 10°85 3-09 1:73 0-72
CaO 10-12 9-80 5-88 4-36 2-53
Na,O 0-54 1-51 2-03 3-14 2°88
K.,0 0-69 0-92 1-03 2-12 2-44
H,O + 0-58 0:65 1-17 1-03 0-50
H,O— 0-13 0:06 0-10 0-06 0:06
TiO. 0:96 1-10 1-42 0-41 0:45
P05 0:30 0-43 0:33 0-19 0-25
MnO 0-14 0-11 0-09 0:08 0-08
Ete = = = 0-32 —
Total th & 100-74 100-59 99-64 100-01 99-59
SpiiGra lon. BA 2-989 2-932 2-768 2-729 2-688

I. Gabbro, Kelly’s Point, 8 miles south-east of Moruya.
II. MDiorite-gabbro, Kelly’s Point, 8 miles south-east of Moruya.
Till. Tonalite, Kelly’s Point, 8 miles south-east of Moruya.
IV. Granodiorite, 2 miles east of Moruya.
V. SBiotite-granite, 3 miles north-west of Moruya.
Analyst, I. A. Brown, Proc. LINN. Soc. N.S.W., liii, 1928, p. 182.

accessory minerals, apatite and magnetite. Olivine is sometimes present. Some
varieties are amygdaloidal, the vesicles containing chlorite and calcite or even
zeolites. Certain phases have suffered deuteric alteration in some localities (W. R.
Browne and H. P. White, 1928).

(ii). In the Milton District, monzonitic rocks occur as intrusions in
horizontally-bedded Upper Marine sandstones which are stratigraphically below the
tuffs of the Illawarra District.

The petrology of the Milton district has been described by G. W. Card (1905,
1915) and subsequently by the writer (19250). The main intrusion outcrops over
an area of about 16 square miles and has a thickness probably not exceeding 400
feet. Sill-like intrusions of similar rocks occur at Little Forest, the Pointer
Mountain and Yatteyattah, which vary from 50 feet to 300 feet in thickness. The
principal rock-types are monzonite, monzonite-porphyry and banatite, the latter
being a fine-grained marginal phase of the intrusion. Some of the more coarsely
crystalline phases have suffered deuteric alteration.

A somewhat similar intrusion occurs at Woodburn, 15 miles south of Milton,
and a more coarsely-crystalline phase outcrops on the coast near Murramarang and
Bawley Point, apparently as an interbedded flow in the Upper Marine sandstones.
The latter includes a type of olivine-monzonite approaching an essexite (L. F.
Harper, 1915, p. 310; I. A. Brown, 1925), p. 464).

BY IDA A. BROWN. 345

(iii). The igneous complex at Mt. Dromedary, 70 miles south of Milton, has
been described in detail by the writer (19300). It outcrops over an area of about
25 square miles, including the whole of the Mountain. The distribution of the
rock-types comprising the intrusion indicates that it is in the form of a laccolith
at least 3,000 feet in thickness. The original cover has been worn away entirely,
and the surrounding rocks consist of highly folded older Palaeozoic sediments. A
study of the occurrence of Upper Devonian sediments along the South Coast
suggests, however, that the Upper Marine stage formerly extended over the area
now occupied by the Dromedary Mountain (I. A. Brown, 1931). Since the Upper
Devonian rest on Ordovician sediments with a very strong unconformity, it seems
quite possible that the igneous injection took place along this unconformity, and
that the intrusion was in the form of an inter-formational laccolith.

The intrusion is essentially monzonitic in character and includes a great
variety of plutonic and hypabyssal types. The main plutonic series consists of
banatite, monzonite, olivine-monzonite and pyroxenite, with a related series of
nepheline-bearing rocks, ijolite, nepheline-monzonite and nepheline-shonkinite or
covite, and a series of melanite-bearing pyroxenites and melteigite-jacupirangites.
Associated with these are some hypabyssal monzonitic rocks which resemble some
of the Milton types. These occur in the form of minor plug-like intrusions about
the foothills of the Mountain, and contain fragments of the more basic monzonitic
rocks and also essexite and other types not known to be exposed in the main
intrusion. Similar rocks outcrop on Montague Island, seven miles east of Narooma.

Table 5 illustrates the chemical composition of the Permian monzonitic rocks;
the analysis quoted in column i is representative of the latitic flows of the
Illawarra, that in column ii of the monzonite-porphyry at Milton, and the remaining
analyses are of the chief plutonic types in the monzonitie series at Mt. Dromedary.

(d). Tertiary.

Associated with the Kamilaroi and Triassic sediments of the Illawarra District
are a number of alkaline rocks and rocks with alkaline affinities, which occur as
dykes, sills, voleanic necks and volcanic flows.

These rocks have been described in considerable detail by L. F. Harper (1915),
G. W. Card (1915), and others. The variety of comparatively rare rock-types
developed in this area makes it one of great petrological interest. The rocks
comprise (i) several varieties of lamprophyre, including mica-lamprophyre,
hornblende-lamprophyre or camptonite and a number of monchiquites; (ii) syenite,
nepheline-syenite and tinguaite; and (iii) olivine-bearing and analcite-bearing
basalts and dolerites.

This series is believed to be of Tertiary age, but the exact age of particular
members can rarely be determined. By analogy with igneous rocks of similar
character in other parts of the State it is generally considered that the lampro-
phyres and alkaline syenites are of early Tertiary age, possibly Hocene, and that
the basaltic rocks are somewhat younger. Some of the latter overlie sediments
containing Tertiary plant-remains. T. G. Taylor and D. Mawson (1903, p. 346)
were also of the opinion that the basalt flows of the Bowral district were sub-
sequent to the syenitic intrusions, and were probably of late Tertiary, Pliocene, age.

(i). The lamprophyres occur as volcanic necks in the Good Dog Mountains
and near Robertson (Wallaya), and also as dykes. The variety monchiquite,
consisting of augite, magnetite and analcite, occurs as sills at Mt. Nebo (west of
Wollongong) and at Rixon’s Pass (west of Bulli), and as numerous dykes (G. W.

346 GEOLOGY OF THE SOUTH COAST OF NEW SOUTH WALES,

Card, 1915, p. 358). The lamprophyres are always porphyritic, the phenocrysts
being pyroxene, amphibole or mica, set in a very fine-grained groundmass composed
of some of the following minerals: plagioclase, orthoclase, pyroxene, hornblende,
quartz, apatite, sphene and iron ores.

TABLE 5.
I II III IV. Vv VI Vil

SiO, 52-42 53:21 43°63 48°34 51-14 59°44 64-49
Al,O; 18:05 17-84 7°52 - 11:79 16-91 19-58 17-48
Fe,03 4-30 3°80 6°45 2°31 1:34 0-31 1:64
FeO 3°60 5:22 8°57 7°72 7:27 3°91 1-69
MgO 3°60 2:96 13-67 9-59 5°88 1-27 0-66
CaO 6-14 6:48 17°12 12°76 9-68 3°95 3°28
Na,O 3°75 3°36 0:36 1-60 1°92 3°21 4-16
K,0 4:14 3°03 0-50 3°17 3°32 6-60 4-79
H.O+ 1:47 1:27 0:46 0-68 0-54 0-88 0:52
H,0— 1:07 0:65 0:25 0-04 0-20 0-12 0:18
TiO. 1:16 1:01 1:24 0-88 0-92 0-54 0-46
P.O; 0:34 0-44 tr. 0-87 0°53 0:07 0-22
MnO 0:28 0:32 0-20 O15 0-14 0-07 0-11
Etc. 0-28 0:29 = = = 0-49 0-77

Total ae 66 100-60 99-88 99-97 99-90 99-79 100°44 100-45

Sim Che oc 0.0 2°722 2-768 3°393 3°085 3°017 2-679 2-653

I. Bumbo Flow, Bumbo Quarry. Anal. H. P. White, Rec. Geol. Surv. N. S. Wales,
viii, 1905-1909, p. 12.

II. Latite (Banakite), Milton. Anal. A. H. Greig, Mem. Geol. Surv. N. S. Wales,
Geology 7, 1915, p. 320.

III. Pyroxenite, Tilba Tilba Lake. Anal. I. A. Brown, Proc. LINN. Soc. N.S.W.,
lv, 1930, p. 664.

IV. Shonkinite, Tilba Tilba. Anal. I. A. Brown, Proc. LINN. Soc. N.S.W., lv, 1930,
p. 656.

V. Olivine-Monzonite, two and a half miles south-west of Tilba Tilba. Anal. I. A.
Brown, Proc. Linn. Soc. N.S.W., lv, 1930, p. 653.

VI. Monzonite, north-west of Tilba Tilba. Anal. I. A. Brown, Proc. Linn. Soc.
N.S.W., lv, 1930, p. 647.

VII. Banatite, Mt. Dromedary. Anal. I. A. Brown, Proc. Linn. Soc. N.S.W., ly,
1930, p. 644.

Several of the lamprophyric and monchiquitic dykes contain large xenocrysts
of brown hornblende and augite, and ultrabasic xenoliths in monchiquitic dykes
are recorded by W. N. Benson (1914) and C. A. Sussmilch (1905). A lamprophyric
dyke south of Moruya (I. A. Brown, 1929) contains xenoliths of ultrabasic plutonic
rocks, lherzolite and pleonaste-pyroxenite, and numerous xenocrysts of red garnet,
brown hornblende, augite and basic plagioclase.

Chemical analyses of twelve lamprophyres and monchiquites have been carried
out in the laboratory of the Department of Mines, Sydney, and are recorded by
L. F. Harper (1915, pp. 325-360). A few typical analyses are quoted in Table 6.

BY IDA A. BROWN. 347

TABLE 6.
I II II. IV Vv

SOA 96 aie ake 46°50 51-02 39°35 41:72 42°52
Ow. 16°48 16°98 11-60 16°87 16°82
Fe,05 3°80 5-52 3-32 5-90 2-90
FeO 7°47 4:13 7°38 3°87 7 +56
MgO 3-46 4°13 9-82 4-23 6-43
Cao 6-14 6-37 12-66 10°32 9°78
Na,O 4°19 3-61 2-00 4:99 4-02
K,0 4°54 2-98 1:77 2-73 1°3
H,O+ 3°57 1-95 3-82 3-67 5:51
H,0— 0-21 1-01 1-28 0-45 0-47
CO, 0-28 0°35 4-04 2-54 0-07
TiO» 2-40 1-44 1:98 1:70 1-30
P.O; 1:13 0-30 0-64 0-85 1-46
MnO 0-15 0-29 0-21 0-15 0-15
BaO 0-08 0-25 0-08 0-13 0-04
Etc. tr 0-28 0-55 0-12 0-03

MAAN ngon le 100-40 100-61 100-50 100-24 100-37

SpeGranen as 2-765 2-788 2-915 2-732 2-812

I. Mica-lamprophyre, volcanic neck (?), Robertson. Anal. H. P. White, Mem. Geol.
Surv. N.S.W., Geol. No. 7, 1915, p. 338.

Il. Hornblende-lamprophyre, dyke, Good Dog Mts. Anal. J. C. H. Mingaye, ibid.,
328.

III. Monchiquite, Kiama. Anal. H. P. White, ibid., p. 360.

IV. Monchiquite (Fourchite), Mt. Nebo. Anal. J. C. H. Mingaye, ibid., p. 344.

V. Monchiquite, Rixon’s Pass. Anal. H. P. White, ibid., p. 348.

(ii). Alkaline syenites also occur as sills and volcanic necks on the Illawarra
Coast and adjacent tableland. The Jamberoo sills consist of nepheline-syenite and
the Dhruwalgha sill is of finer-grained tinguaite. With these may be grouped the
volcanic necks of the Bowral Gib and Mt. Jellore, described by T. G. Taylor and
D. Mawson (1903) and which consist essentially of alkaline syenite associated with
alkaline basic and ultrabasic rocks, essexite and picrite.

Chemical analyses of the principal types are quoted in Table 7.

(iii). The basalts and dolerites occur as dykes, sills and volcanic flows. The
dykes are most easily identified along the coastal rock-platforms and usually consist
of fine-grained basaltic rock. The sills show greater variation in texture and
composition although they are not of great thickness or extent. Basaltic flows form
a .capping to the Illawarra Tableland in the vicinity of Robertson, and south of
the Shoalhaven River basalt occurs on the Sassafras tableland (G. W. Card, 1915,
p. 286); thin flows outcrop near Milton, Moruya, Bodalla, Yellow Pinch (Wolumla)
and elsewhere (I. A. Brown, 1925c, 1931). The columnar basalt of Mt. Darragh
is portion of an extensive basalt flow or series of flows, which form the surface
of the Monaro tableland and extend to Bombala and Nimmitabel.

348 GEOLOGY OF THE SOUTH COAST OF NEW SOUTH WALES,

TABLE 7.
if II. Til. IV.

SiO, As “iN ai3 54-50 55-82 95 +86 55-16
AOS ted ae 17-81, 20-19 15-25 16-67
Fe,0; 1-70 Beard) 4:92 230
FeO 5:30 iLoaly/ 6-07 7-31
MgO 1-09 0-25 0-20 0-56
CaO 2-64 1:02 Dealts 2-30
Na,O 5-39 9-57 2-34 5-65
K,O 6-08 5-60 9-28 6:97
H,O+ BiO7KS} 1:30 0-50 0-88
H,O— 0-54 0-15 0-70 0-85
Co, 0:30 0-04 1-80 1-50
TiO, 0-44 0-01 0-65 0-60
P.O; 0:44 0:07 0-16 0-38
MnO 0:25 0:36 0-42 0:47
BaO abs. tr. tr. tr.
Etc. 0-02 0-55 0-14 0-42

Total a ays 100-28 99-80 100:39 100-80

SpaiGLaer. wee 2-595 2-594 2-706 2-675

I. Nepheline-syenite, sill, Kangaroo Valley, Jamberoo. Anal. W. A. Greig, Mem.

Geol. Surv. N.S.W., Geology No. 7, 1915, p. 339.
II. Tinguaite, sill, Dhruwalgha. Anal. J. C. H. Mingaye, ibid., p. 341.
III. Syenite (melanocratic), Bowral. Anal. T. G. Taylor and D. Mawson, Journ.

HOYS SOC INES {Wis EEXXVLl O03 5) py odie
IV. Syenite (leucocratic), Bowral. Anal. T. G. Taylor and D. Mawson, ibid., xxxvii,

19103; 0D. oad.

The basalts are generally compact, and, unlike the Kamilaroi basaltic rocks
(latite), are not usually porphyritiec in felspar. They consist essentially of laths
of plagioclase and augite, the latter mineral occurring either as small rounded
grains producing intergranular fabric, or as plates including laths of plagioclase
and giving an ophitic fabric to the rock. As a rule olivine and iron ores are
present and analcite frequently occurs. Nepheline is present in some cases, as in
the theralitic basalt from Bombala (W. R. Browne, 1927, p. 377), but generally
occurs only in the norm. Table 8 illustrates the chemical composition of these
rocks.

The first four analyses (in Table 8) of New South Wales rocks are remarkably
similar considering that they come from widely separated localities. They are all
undersaturated in silica and show about 20 per cent. of olivine and a small amount
of nepheline in their norms. Three fall into the same subrang, Auvergnose, in the
American Quantitative Classification, while the Cordeaux analcite-dolerite belongs
to the closely related subrang Camptonose.

It is thus evident that these basalts do not strictly belong to the ‘plateau
basalts” in the sense used by H. S. Washington (1922, p. 797), who states: “In the
typical plateau basalts ... . no nephelite is .present, although some nephelite

BY IDA A. BROWN. 349

basalts or tephrites accompany the normal basalts in a few places. .... Olivine is
generally rare,.... Quartz is not usually present in these basalts, although many
of them show an excess of silica in the norm.’ An average of 11 analyses made
by Washington of the Deccan basalts is quoted for comparison in column vi.

TABLE 8.
il Il. III. IV. Vv. VI.

SiO, 44-57 46°20 44°35 44-08 15-25 50°61
Al,O, 15-30 17°44 14:71 16:70 16-47 3-58
Fe.0; 3-20 1:30 3°08 2-77 2°44 3-19
FeO 7°83 9-54 8-21 8-08 8-69 9-92
MgO 10-04 7:38 10-63 9-25 7-77 5-46
CaO .. 10-00 9-02 10:08 9-49 8-90 9-45
Na,O 1:94 3-69 1:64 2-01 2-68 2-60
K,0 1-39 1-10 1-39 1-74 1-42 0-72
H,0+ 3:21 2-11 3:04 2-45 1:60 } 16
H,0— 1:09 0:09 0-72 0-64 0-70 nea
co, 0-01 0:04 tr. tr. —
TiO, 1:01 1:30 1:95 2-03 3-20 1-91
P.O; 0-41 0:56 0:59 0:50 0:43 0-39
MnO 0-29 0:17 0-15 0-16 0-38 0:16
BaO 0-05 0-03 — -- —
Etc. 0-09 — — a — —

Motalue we 100-43 99-97 100°52 99-90 99-93 100-12

Sp. Gr. .. 2-907 2-934

I. Analcite-basalt (Robertson Flow), 5 miles west of Jamberoo. Anal. H. P. White,
Mem. Geol. Surv. N.S.W., Geol. No. 7, 1915, p. 288.

II. Ophitie analcite-dolerite (Cordeaux Flow), half a mile east of Wanyambilli.
Anal. H. P. White, ibid., 1915, p. 289.

Ill. Basalt, 12 miles north of Bombala. Anal. I. A. Brown.

IV. Basalt, half a mile south of Nimmitabel. Anal. I. A. Brown.

V. Olivine-basalt, Lago Buenos Airos, Argentina. Anal. Herdman, Jowrnal of
Geology, xl, 1932, p. 379.

VI. Average of 11 analyses of Deccan basalts. H. S. Washington, Bull. Geol. Soc.
Amer., XXxiii, 1922, p. 797.

On the other hand the basalts are very similar in all respects to the Patagonian
basalt, whose analysis is quoted in column y. This rock is described by G. W.
Tyrrell (1932, p. 374) as a “plateau-basalt in the restricted sense of Gregory and
Reck. This type contrasts with that of flood-basalts, which, in general, are over-
saturated with silica.”

The alkaline affinities of many of the Tertiary basalts of Eastern Australia are
recognized by Sir T. W. E. David (1932, p. 103).

4. Tectonic History AND IGNEOUS ACTIVITY.

The distribution of outcrops and the variation in the conditions of sedimen-
tation as revealed by the lithological and palaeontological characters of the lower
and middle Palaeozoic rocks of the South Coast of New South Wales supplies a

B

350 GEOLOGY OF THE SOUTH COAST OF NEW SOUTH WALES,

considerable amount of evidence on the history of the building of south-eastern
Australia and the changes in the palaecgeography of this region during lower
Palaeozoic time.

The oldest sediments, possibly of Cambrian age, indicate deposition under
marine conditions; normal sedimentation was accompanied by deposition of tuff
and other products of submarine igneous activity. The vast thickness and extent of
this formation suggests that the landmass from which these sediments were
derived was not far distant, and probably was situated to the east. This land may
have been the predecessor of the “Tasmantis” of Stissmilch and David (1919).

The palaeogeography of Australia during the previous Pre-Cambrian period
has been considered recently by L. A. Cotton (1930), who suggests for the Pre-
Cambrian framework (1930, figure 4, p. 55) a main south-western block named
Yilgarnia separated from three borderland masses, the Kimberley massif, the
Carpentaria massif and the eastern massif of Tasmantis, by the Nullagine
geosyncline.

This conception of the Australian framework is comparable with that envisaged
by C. Schuchert (1923) for the continent of North America. Schuchert states
(p. 158): “From the geographic position of the geosynclines near the margins of
the continent, and the further fact that the main masses of their sediments came
not from the medial area of the continent, but from more or less narrow lands
facing the oceans, it is clear that North America was originally much more
extensive than now. Since these facts are already true in Lower Cambrian time,
it is also clear that the extent of greater North America was estabiished in
Proterozoic time.’ These ideas may be applied with equal truth to the continent
of Australia.

The early Palaeozoic history of south-eastern Australia is chiefly that of the
infilling of the southern part of the Nullagine geosyncline. Thus Cambrian (7),
Upper Ordovician, and Upper Silurian sediments were deposited in the New
South Wales portion of the geosyncline, the apparent absence of Lower Ordovician
and Lower Silurian sediments being due either to lack of knowledge of outcrops
or to the possibility that during these epochs the sea was withdrawn to the south
and that no sedimentation took place in New South Wales. Upper Ordovician rocks
extend eastwards to a few miles east of Quaama and Cobargo, the Upper Silurian
to the upper part of the Deua River. The geosyncline during Lower and Middle
Devonian time apparently was narrower than it had been formerly, but extended
from eastern Victoria into New South Wales through Yass and Tarago to the
Mudgee district. During the Lower and Middle Devonian epochs the present South
Coast was an area of non-deposition, probably forming part of the borderland of
Tasmantis.

At the end of the Middle Devonian the southern part of the Nullagine geosyn-
celine was finally closed up and the south coastal portion of New South Wales was
permanently welded on to the continental mass of Australia. As a result of this
diastrophism the Middle Devonian sediments were folded in a meridional direction
and possibly epeirogenic uplift accompanied the orogenic movements (I. A. Brown,
1932).

The Upper Devonian commenced with an epoch of igneous activity represented
by enormous flows of acid volcanic rocks, which form the Eden stage of the series.
No volcanic centres have been recognized; possibly the flows were associated with
fracture and faulting preceding the formation of a marginal geosynclinal trough in

BY IDA A. BROWN. 351

which later sedimentation occurred. Gradual subsidence followed and lacustrine
sediments of the middle or Yalwal stage are associated with contemporaneous
flows of rhyolite and basalts allied to spilites. The upper or Lambie stage of
marine sediments was deposited over a wide area of New South Wales, the deepest
part being towards the east. The western portion of this sea was not geosynclinal,
but was rather of the nature of a flood over the relatively stable continental
massif. Thus a thin veneer of late Upper Devonian arenaceous sediments was
deposited over highly folded pre-Devonian rocks, and these were not afterwards
subjected to intense folding.

The intensity of the diastrophism at the close of the Middle Devonian is
further emphasized by the marked differences in the lithological characters of the
Middle and Upper Devonian sediments, in the deposition of thick clastic sediments
in the Upper Series, and also by the marked palaeontological break between the
two series. Unconformable contact has been observed only in Victoria.

After the deposition of the Upper Devonian sediments orogenic movements
affected the greater part of south-eastern Australia: the Eden district became
portion of the stable land block, which has suffered no later orogenic movement,
and the area of instability moved northwards. This late Devonian orogeny was
aecompanied by the intrusion of great granodioritic batholiths, an association of
fold-movement and intrusion of subalkaline magma which supports the views of
A. Harker (1909) regarding the distribution and origin of igneous rocks.

The elongation of the main batholith described earlier in this paper, and also
the longer axes of the smaller batholiths are parallel to the general direction of the
trend lines of the Upper Devonian sedimentary rocks of the South Coast.

There are no signs of Carboniferous sedimentation in the area, and it is
inferred that the area formed part of a land-mass during this time.

The development of the Kamilaroi geosyncline to the north and east resulted
in heavy sedimentation. During the Upper Marine stage there were vertical adjust-
ments of the adjacent landmass (without strong folding movements) and these
were accompanied by the deposition of tuffs and flows of latite in the Illawarra
district, by sill-like intrusions in the Milton district and by laccolithic intrusion
in the Mt. Dromedary district. These intrusions were monzonitic in character, or
rather more alkaline than those of the preceding period of igneous activity.

No important orogenic movement has taken place on the South Coast since
the close of Palaeozoic time. The axis of geosynclinal sedimentation gradually
moved northwards towards Sydney and post-Palaeozoic earth-movements were
mainly vertical adjustments, broad warping with some faulting. These movements
were accompanied by the extravasation of highly differentiated alkaline magma in
the form of dykes, sills and volcanic necks. The earlier intrusions were lampro-
phyric and syenitic, and the later occurrences were of alkaline basalt.

The association of intrusions of more alkaline rocks with the later epeirogenic
movements of the area again supports the generalization of A. Harker (1909)
concerning the relation of the chemical composition of igneous intrusions to the
associated type of earth-movement.

The Tertiary earth-movements affected the whole of eastern New South Wales
and culminated in general elevation during the Kosciusko epoch. The east-west
trend of the majority of the dykes on the South Coast suggests that the crust of the
earth here was in a state of relative tension in a meridional direction. This may be
an expression of relative compression in an east-west direction, for the plateau-

352 GEOLOGY OF THE SOUTH COAST OF NEW SOUTH WALES,

building forces operated in a direction at right-angles to the trend of the coast-line
and the coastal plateaux.

More recent tectonic history is concerned mainly with oscillatory movements
about the present coast-line and the development of the present physiography. No
igneous activity is known to have occurred since Tertiary time.

5. Magmatic DIFFERENTIATION.

Many problems of magmatic differentiation present themselves in a study of
the igneous rocks of the South Coast. These may be divided into two groups:
(a) those of a local character, which are related to the petrogenesis of rock-series
occurring in a limited area, and (0) those of a regional character, which are con-
cerned with the ultimate consanguinity of rock-series of several petrographical
provinces.

(a). The effects of local magmatic differentiation have been considered
previously by the writer in the cases of the Milton (19250), Moruya (1928) and
Mount Dromedary (19300) intrusions, but for the sake of completeness these will
be summarized in the following account.

— Basalt

20

[rea = Granodiorite

e |
oy:
yi
He lee a
a

20

15

| 4g0 ae ee
ate ee

ae

fe
60
S

O
u a S—_ —SS
0 | KO SS Slee ea) iO.
SiO ES 8 S 3 2 2

Text-figure 1.

There can be no doubt that the South Coast was portion of a petrographical
province during the early and middle Palaeozoic era, and that the igneous
intrusions and extrusions were typically subalkaline in character. The widespread
extrusions of rhyolite and basalt during the Devonian suggest that similar
magmatic and tectonic conditions obtained over a zone several hundred miles in
length, following the margin of early Upper Devonian sedimentation. There is
no obvious genetic relationship between the rhyolites and the basalts, but the
granodiorite, which was injected as batholiths at the close of the Devonian, is
possibly related to both the earlier volcanic types.

The accompanying variation-diagram, Text-figure 1 (of the type used by
A. Harker, 1909) is based on three analyses, which are regarded as typical and

BY IDA A. BROWN. 353

representative of the (Devonian) rhyolites, granodiorites and basalts of the South
Coast. These are of (i) the basalt from Nethercote, near Eden (p. 340), (ii) the
granodiorite from Moruya (p. 344) and (iii) the rhyolite from the Lower Deua
River (p. 339) whose analyses have been quoted in an earlier part of the paper.
The points representing the various oxides are joined by simple curves, the form
of the alumina curve towards the basic end being determined by comparison with
the sum of the other oxides.

The resultant diagram is that of a typical subalkaline series, with the
eharacteristic features as defined by Harker (1909, p. 130): “The flat convex shape
of the alumina curve, the declining concave curves of magnesia and ferric oxide,
the lime with its maximum near the basic end, the soda with its maximum near
the acid end, and the steady rise in the potash-line’’.

It is therefore considered by the writer that the Devonian rhyolites and basalts
are genetically related to the granodiorites, and that they represent complementary
differentiates of the granodioritic magma. The cause of the differentiation is not
apparent. Evidently differentiated magma underlay the region during early and
middle Palaeozoic time; the early Devonian earth-movements were probably of
the nature of trough faulting and broad warping, which permitted the extravasation
of the acid and basic differentiates as volcanic flows. An interesting feature
is that, as previously shown (1931), the basalts are allied to spilites and are
associated with movements of subsidence, a feature which is characteristic of
spilites in other parts of the world. The main granodioritic magma was not
injected until after the close of the Upper Devonian, during a period of crustal
folding.

In a recent paper to the Geological Society of London (1932), Dr. A. Brammall
has indicated that possibly the Dartmoor granites and the spilites of Devon and
Cornwall are genetically related. In the discussion following the reading of the
paper, Sir John Flett pointed out that in Ayrshire... “‘the spilitic eruptions had
been foHowed, as in Devon, by a period of intense folding. After a short interval
the folded rocks were invaded by a new magma, represented by the Galloway
granites.” :

The similar magmatic and tectonic conditions during Devonian time in Britain
and on the South Coast of New South Wales, certainly indicate some causal
connection between the processes of magmatic differentiation and earth-movement.

The granodioritic magma injected after the deposition of Upper Devonian
sediments on the South Coast took the form of a large batholith, extending from
Victoria into New South Wales, and several smaller intrusions, such as those near
Bodalla, Moruya, Coondella and Nelligen, which may be regarded as apophyses of
the main batholith.

Although the writer has not yet had the opportunity of carrying out a
chemical investigation of the main batholith, its field and petrological examination
indicates that differentiation tock place in a manner similar to that which occurred
at Moruya. It has been shown (1928) that here the igneous rocks form a complete
and typical subalkaline complex, as indicated by the accompanying variation-
diagram (Text-figure 2).

The differentiation was of a serial character, and probably took place by
means of fractional crystallization and the sinking of crystals in the manner
postulated by N. L. Bowen (1915, 1919). It was considered (1928, p. 186) that
this process took place in an intercrustal magma-chamber and that the apparent
intrusive relationships between the chief plutonic types were due to successive

354 GEOLOGY OF THE SOUTH COAST OF NEW SOUTH WALES,

injections of magma of increasing acidity and increasing alkalinity from the
hypothetical magma-reservoir. There is no clear evidence to show that such has
been the case, and the increasing number of described occurrences of a similar
nature in which magmatic differentiation has occurred practically in place, suggests
that a similar process may have taken place in the Moruya magma, and that slight
earth-movements during its consolidation are responsible for the apparent intrusive
relationships.

° 3 e
= ~ iS >
= = vee = 3
SS “A hata nd a i~d
5 S sae es
o 2 ~ LSS Q =
©) & aS = sh s YX
co c= Pay ~ SSR = % 2
as = 3 ZES Ss =
@ cs S° = ris 3 5 a
S =P =
SA a ‘Ss 5 OES SS
] T
ah | j a lenis
‘clmaleuealie oH Tapa Haale Tlaeastl
e 9 ty 1 1 1
Hilt [! | I

Text-figure 2.—Variation diagram for the igneous complex of the Moruya district.

Magmatie differentiation may have proceeded as well in the large apophyses as
in the main batholith, without any intermediate magma reservoir as such, and with
the production of similar rock-series on different scales of magnitude. Possibly the
greater volume of magma in the main batholith and the resultant slower cooling
is responsible for the formation of more coarsely-crystalline rocks than the
mineralogically equivalent types in the smaller intrusions. The formation of
complementary dyke-rocks, aplites and dolerites, took place during the later stages
of consolidation of the magma. Associated with these and the final stages are
metalliferous deposits, gold and ores of bismuth, molybdenum, arsenic with traces
of silver, lead, zine and tin.

12 vs
1 a ;
10 1 wa | el nr i
9 L He + Hue
’ ' 7 1 \ |
8 : i nia a sail
Cf = aL a niles ; joel
1 vets 1 u |
6 —— : : at i tts aa (mea aaa (a
5 Ke He ff | ‘ent tlie | |
: ! a; eae | 30 ee
: Sse if aa |
Fa : : i a : :
2) la = aL — g Cog. "
TiO, ! . A {
i = = 2 : =<
KO Tio z = Meg
1/3, Ww) S
Silica—> ~ 35 os Ss Sos S55 5 RX x i

&0

ceo
ol
ol

BY IDA A. BROWN.

The flows of the Illawarra District give evidence of slight progressive magmatic
differentiation during late Kamilaroi time, but this variation is only of the order
which obtains within one of the larger flows, such as that of Bumbo (G. W. Card,
1915, p. 285). Late-magmatic or deuteric processes have operated to a considerable
extent in some of these flows, as shown by the work of W. R. Browne and H. P.
White (1928).

Somewhat similar conditions prevail among the Milton monzonitic rocks (1. A.
Brown, 1925b). There are only slight variations in the chemical composition of
different phases of the main intrusion, although there are differences in texture
and mineralogical composition. The occurrence of cognate xenoliths and aplitic
and pegmatitic segregation veins indicates greater magmatic differentiation than
appears to have taken place among the volcanic equivalents of the Illawarra.
Possibly this is due to the form of the intrusion and the depth of cover at the time
of injection, which permitted slower cooling and crystallization. During the final
stages of consolidation, albitization and associated processes operated in this mass
also, particularly in the more coarsely crystalline portions.

The igneous complex at Mount Dromedary gives evidence of much more
complicated differentiation than has taken place in the other monzonitic occur-
rences of the South Coast (I. A. Brown, 19300, pp. 688-691). The intrusion is in
the form of a laccolith which probably had a fairly thick sedimentary cover at the
time of injection. The variety and arrangement of the rock-types developed in this
occurrence indicate that the relatively great thickness (3,000 feet) and slow
cooling of the laccolith afforded opportunity for the differentiation of the magma
in situ, by means of fractional crystallization and the sinking of crystals under
gravity, as postulated by Bowen (1915, 1919), Harker (1909, p. 317; 1913) and
others.

As a result, the main monzonitic series, including the banatite, monzonites,
shonkinite and pyroxenite, was produced. Probably earth-movements during the
consolidation of the magma were responsible for the partial injection of the
banatite into the monzonite, and also indirectly for the development of an
associated series of nepheline-bearing rocks, nepheline-shonkinite and nepheline-
monzonites, as a result of filter-press action. The origin of the associated garnet-
bearing series is obscure. The accompanying variation-diagram (Text-figure 3)
illustrating the chemical composition of the occurrence indicates that the garnet-
bearing rocks are not directly related to the monzonitic series, although field and
mineralogical evidence is in favour of some degree of consanguinity.

The possibility that the garnet-bearing rocks are due to the incorporation of
limestone by the monzonitic magma has been considered previously by the writer
(1930b, pp. 690, 691), and no further information is available on the subject.

The later stages of consolidation are represented by aplitic and lamprophyric
dykes and veins in the main intrusion. The formation of volcanic necks or plugs,
containing inclusions of the plutonic types, marks the final phase of igneous
activity in this locality.

The igneous rocks which have been referred to the Tertiary period show great
diversity, which is characteristic of alkaline rocks all over the world.

Magmatic differentiation evidently took place in an intercrustal reservoir and
successive periods of injection are indicated by numbers of small intrusions, which
are more or less homogeneous in composition and which may be grouped into
series possessing certain distinctive chemical characters. Thus the early intrusions

356 GEOLOGY OF THE SOUTH COAST OF NEW SOUTH WALES,

of lamprophyres are basic, distinguished by relatively high alumina and high
alkalis; the monchiquites are still more basic with lower alumina, but with higher
magnesia, lime and water. Similar variations occur among the nepheline-syenites
and dolerites which were probably injected at a somewhat later period.

The basalts are probably the latest expressions of igneous activity in the area.
These are not normal basalts, but show decidedly alkaline affinities, indicated
mineralogically by the presence of analcite and (normative) nepheline in almost
every case, and shown chemically by the relatively high amounts of alkalis and
titania.

3
=
i=]
2 3 8
18 2 35 2 33 S32
a e ES = 23 3 is 2
x 8 3 2 R= 8 =£§ =
Ss 3 = gS bu =
St ae GE ss 3 : &
s S a iS _ N
'
i: d :
' U 1 1 "

20 ; rr = ALO, : H
H 6 ' 26 q ' H

19 ‘ ote ' H
yt , i |

wo 1 Bally ' b ' '
‘ tye ' t ! °

17 ' r
i : ' 1 1
q H ' ; i

Ls) cy uml H q i '

N a ' d } 4 4 '

ae aly ale: ils |
4 NO ' H 1 '
<r Peaps
» ; B 1 ' '

13 a Smilin 1 \ 2
H NX) SN! ! |

12 2 Tv 1 '

N A aTAy Qa ’

7 ' os Sh Hall <o i '
= ay ’ i
fe Pal Se !

OR 5 );
: ; ! ' ' q

8 1 alee 1 H
ies |
® } Vaal BT Nee:
' ' it ' '

? 4 i r
“aa 4 20
5 Ol Ale 0 f !
, Bld ! qi | t
0,’ tt ! 1 | :

8

¥ H K,0 > Bolt 3

A \

‘ 1 MgO ! f i)
‘ ' ' Na,0 i < +
3
i) i ' 1 i 1 '
pleeal me tee eo
tela | ” } ' i
I er Fai eo : Tio 3
as
ia as O) ! ' #0, ° = 9
S
ES Sma 8 Ss a GSS = S Ss ps8

Text-figure 3.

Differentiation in situ oecurred only in the larger intrusions, such as those of
the Good Dog Mountains and Bowyral.

(b). The possibility that magmatic differentiation has operated over a large
region throughout the whole of geological time, is suggested by the progressive
change in alkalinity and basicity of the injected magma in the area under con-
sideration. Tectonic disturbances at various periods have provided a means for
the injection of magma into the upper layers of the earth’s crust, and at times may
have assisted the process of magmatic differentiation by filter-press action, the
squeezing out of the liquid portion from a partially solid mass. Magmatic

BY IDA A. BROWN. 357

differentiation has been intensified shortly before and sometimes after diastrophic
periods when igneous activity has occurred.

Thus during the middle Palaeozoic the South Coast was essentially a sub-
alkaline province, but complementary differentiates, which formed rhyolites and
basalts, were extravasated before the injection of the granodioritic magma, and
probably differentiation of the latter took place almost in sitw in the batholiths.

In Kamilaroi time the average composition of the magma was monzonitic, more
alkaline and more basic than that of earlier time, and the magma was not highly
differentiated before injection to its final position, but under suitable conditions,
as at Mt. Dromedary, magmatic differentiation was able to proceed almost to its
possible limit, and a great variety of monzonitic and associated alkaline types were
produced.

In Tertiary time the average composition of the mother-magma was that of an
alkaline basalt, the actual weight percentage of alkalis being less than those of
the average monzonitic or subalkaline series, but relatively high for a basic
magma. Here again, intensified magmatic differentiation prior to extravasation
was the general rule, for individual intrusions are of fairly uniform composition
and only under particularly favourable conditions did differentiation occur in the
final position of consolidation.

The change in average composition of the magma injected during these three
epochs is illustrated in the accompanying variation-diagram (Text-figure 4), based
on the analyses of (i) the Moruya granodiorite (Devonian), (ii) the Milton
monzonite (Kamilaroi) and (iii) the Robertson flow of alkaline basalt (Tertiary),
which are regarded as the most representative available analyses.

The resultant curves for the various oxides give a diagram, which is typical
of a subalkaline igneous series, and which may be compared directly with the
variation-diagram for the Moruya complex (Text-figure 2) for a similar range of
the silica percentage. It is considered that this detailed similarity indicates that
the rocks (or magmas they represent) are genetically related, and that therefore
the subalkaline, monzonitic and alkaline series of the South Coast are
consanguineous.

If this conclusion be justified, then magmatic differentiation has operated on
(potential) magma over a long period of time, at least from the Devonian period
to the late Tertiary. It is not intended to imply that this magma was in a fluid
state throughout the whole of geological time, but that diastrophic periods have
afforded opportunity for the liquefaction and injection of portions of the magma
into the upper layers of the earth’s crust, where subsequent magmatic
differentiation, influenced by local factors, has preduced various series of related
rocks.

The Tertiary alkaline rocks of the South Coast are regarded therefore as
normal products of differentiation: the writer is not aware of any evidence that
their occurrence is influenced by association with limestone, as implied by R. A.
Daly (1910, 1914, Appendix D, p. 523), for the intruded Kamilaroi and Triassic
sediments contain no massive limestones, and the underlying Upper Devonian and
older Palaeozoic series are quite devoid of limestone in this region.

It is considered by the writer that further evidence of the production of
the chief occurrences of alkaline rocks as a result of normal magmatic differen-
tiation over a long period of time is afforded by the conclusion of H. I. Jensen
(1908, p. 585) that “Almost all the great alkaline eruptions of all parts of the earth

358 GEOLOGY OF THE SOUTH COAST OF NEW SOUTH WALES,

took place in the Hocene period” and that these occurred “along the borders of
Mesozoic epi-continental basins or transgressions which have been broadly uplifted
without much folding”.

I Ul

_ “-
15

TERTIARY
PERMIAN
DEVONIA

Text-figure 4.

Monzonitic series in other parts of the world are frequently younger than the
subalkaline rocks with which they are associated: in most cases where their age
can be proved it is found to be late Palaeozoic or Mesozoic, that is, in general,
intermediate in age between the subalkaline and alkaline series of any particular
region.

It therefore seems possible that the origin and relationships of the igneous
rocks of the South Coast of New South Wales may be typical and representative
of conditions in other parts of the world, and that in any particular region, which
contains a record of geological history throughout several eras, magmatic differen-
tiation has operated on a grand scale throughout geological time. The earlier
intrusions are subalkaline in character and the later ones are progressively more
basie and relatively more aikaline: their manifestation at the surface of the earth
has been brought about by tectonic disturbances, which may sometimes have had
an influence on the chemical composition of the injected magma. The subsequent
differentiation of a particular mass of injected magma has been dependent on local
conditions, and various factors have operated to produce a great variety of igneous
rocks.

6. SuMMARY AND CONCLUSIONS.

The South Coast of New South Wales may be regarded as a section across the
margin of a continental mass, which has been growing by the repeated deposition
of sediments along its borders and their subsequent elevation and compression to
form portion of the land-mass.

BY IDA A. BROWN. 359

The paper gives an account of the distribution, lithological characters, mutual
relationships and tectonic structures of the sedimentary rocks, whose ages range
from Cambrian (?) to Post-Tertiary, and summarizes the field-occurrence and
petrological and chemical characters of the associated igneous rocks, which belong
to three principal periods of igneous activity. These are referred to the Devonian,
Kamilaroi and Tertiary periods, which are represented by rocks of subalkaline,
monzonitie and alkaline facies, respectively.

A consideration of the tectonic history shows that during early Palaeozoic time
the present south coastal area was portion of a “borderland”, which was separated
from the continental massif by the Nullagine geosyncline, an inheritance from
Pre-Cambrian time. The early Palaeozoic history of south-eastern Australia was
essentially that of the infilling of this geosyncline until the close of the Middle
Devonian, when, during a period of diastrophism, the present south coastal district
became portion of the continental mass. This remained a positive area during the
Carboniferous. The area to the north and east gradually subsided and developed
into the geosyncline of Kamilaroi and Triassic times, in which the axis of greatest
sedimentation moved slowly to the north and east. Cainozoic earth-movements
were chiefly epeirogenic.

There is a close relationship between the history of igneous activity and the
tectonic history of the region. Terrestrial flows of rhyolite and basalt heralded
subsidence and the deposition of Upper Devonian sediments; the subsequent
folding of these beds was accompanied by the intrusion of granodioritic (sub-
alkaline) batholiths, whose elongation is parallel to the axes of folding of the
Upper Devonian sediments. Vertical adjustments about the borders of the
Kamilaroi geosyncline were associated with intrusions and extrusions of monzonitic
magma during Upper Marine time, and epeirogenic movements during the Tertiary
period were accompanied by the extravasation of basic, alkaline magma.

Thus, viewed broadly, the relations of the subalkaline and alkaline rocks of
the region under consideration to the accompanying orogenic and epeirogenic earth-
movements are essentially in accordance with Harker’s generalization on the
subject.

The progressive change in the basicity and alkalinity of the injected magma in
the region under consideration suggests an ultimately comagmatic origin for all
the igneous rocks, which is confirmed to some extent by the form of the variation-
diagram, based on representative chemical analyses of Devonian (subalkaline),
Kamilaroi (monzonitic) and Tertiary (alkaline) rocks.

It is concluded that probably magmatic differentiation has operated on
potential magma for a long period of time, extending at least from the Devonian to
the Tertiary periods on the South Coast. Diastrophic movements have afforded
opportunity for the liquefaction and injection of portions of the magma into the
upper layers of the earth’s crust, and sometimes may have assisted the process of
differentiation by filter-press action. Magmatic differentiation was intensified
sometimes before, and usually after, injection of the magma into its final position,
where various local factors have influenced the subsequent course of differentiation
and the production of the closely related rock-series which are described in the
paper.

The alkaline rocks are regarded therefore as normal products of magmatic
differentiation, and there appears to be no evidence that they are due to
assimilation of limestone in this region.

360 GEOLOGY OF THE SOUTH COAST OF NEW SOUTH WALES,

A brief general survey of the occurrence of subalkaline and alkaline rocks in
other parts of the world suggests that the origin and relationships of the igneous
rocks of the South Coast may be typical and representative of conditions
elsewhere.

The paper is accompanied by variation-diagrams illustrating local and regional
magmatic differentiation, and by a geological sketch-map of the South Coast of
New South Wales.

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, 1919.—Crystallization Differentiation in Magmas. Journ. Geol., xxxvii, 1919.

BRAMMALL, A., 1932.—The Dartmoor Granites: their Genetic Relationships (with 80
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, 1930a.—The Geology of the South Coast of New South Wales. ii. Devonian
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, 1930b.—Idem. iii. The Monzonitic Complex of the Mount Dromedary
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Browne, W. R., 1914.—The Geology of the Cooma District, N.S.W. Journ. Roy. Soe.
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, 1927.—Petrological Notes on some New South Wales Alkaline Igneous Rocks.
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BY IDA A. BROWN. 361

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362 GEOLOGY OF THE SOUTH COAST OF NEW SOUTH WALES.

WASHINGTON, H. S., 1922.—Deccan Traps and Other Plateau Basalts. Bull. Geol. Soe.
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EXPLANATION OF PLATE XXVIII.
Geological Sketch-map of the South Coast of New South Wales.

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THE COCCIDAE OF THE CASUARINAS.
By Water W. Froaearr.
(Plate xxix; thirty-six Text-figures.)
[Read 27th September, 1933.

The Casuarinas, popularly known as Sheoaks or Bulloaks, are remarkable in
having no true foliage. The main branches spread out from the stem and carry
slender cylindrical jointed branchlets, with each ring encircled with small bracts
which represent the leaves.

As might be expected, the insect fauna of the Casuarina is as interesting as
the tree, for they adapt their structure to the form of the branchlets. The insects
which feed upon, or attach themselves to, the branchlets are usually elongate or
angulate when they affix themselves at the base of the branchlet with its attach-
ment to the stem.

The members of the two gall-making genera, Cylindrococcus and Frenciia,
were first collected in Victoria by French, who sent them to New Zealand for
Maskell to name and describe. The first are wonderfully like the seed cones of
their host trees and may be formed upon the branches or the tip of the branchlets.
The second are double galls, forming on the stems, the basal portion forming a
swelling of the tissue, from which, in the typical form, projects a hollow tube,
rounded at the tip, in which the female coccid is resting.

In the following pages I briefly describe these, list other coccids previously
described, and add some new species.

ASPIDIOTUS CINGULATUS Green.

This is a very distinctive species, which I figured and described in my
Descriptive Catalogue (pt. 1, p. 10). It comes from the Mallee scrublands of
North-west Victoria, infesting the branchlets of Casuarina lepidophloia. The large
dark brown scales are so broad that they curl round the branchlets they infest.

GYMNASPIS SERRATUS, ND. sp. Text-figs. 21, 22.

The female tests black, shining, cireular, convex, scattered all over the
branchlets. Diameter 1 mm. Habitat: New South Wales, on Casuarina sp.

®. Coccid light chocolate-brown, broadly oval, broadest across the posterior
portion. Derm showing no definite hairs or spines, slightly crenulated on the sides.
Antennae composed of eight segments of uniform thickness: 1 very short, 2 small,
3 longest, 4, 6 of uniform length, 7 small, 8 irregular in form; the whole fringed
with fine hairs from segment 3 to the tip. Pygidium with a pair of finely
striated chitinous plates, hardly projecting beyond the margin of the anal segment.
The whole almost truncate, with the inner area between the plates angular.

LEPIDOSAPHES (MYTILASPIS) CASUARINAE Maskell.
This is the most abundant and widely spread coccid infesting the Casuarinas.
Usually the branchlets are so thickly incrusted with the white scales that they
are very noticeable.

364 THE COCCIDAE OF THE CASUARINAS,

The white female test is so finely striated that, in a bright light, it is
iridescent. The pellicles are reddish-brown, the first oval and very prominent;
the second large, rounded behind, and fitting in the test. Length, 2 mm. It is
variable in size, those upon the Myall and native cherry being larger than the
type.

Hab.—Sydney, on Casuarina suberosa; Pilliga Scrub and Euston, N.S.W., on
Casuarina lepidophloia; Leeton, N.S.W., on the same species (Keith McKeown) ;
Millmerran, Queensland, same species (J. Macqueen); Trangie, N.S.W., on
Casuarina Cambagei (W.W.F.); Perth, W.A., Casuarina sp. (L. J. Newman). Also
upon Hxocarpus cupressiformis, Crookwell, N.S.W., and Acacia sp., Dubbo, N.S.W.
(W.W.F.).

LEPIDOSAPHES HILL Laing.

Bull. Entom. Research, xx, pt. 1, 1929, p. 118, f. 17.

This species is closely allied to L. casuarinae; Mr. Laing says that the external
characters of the puparium are very similar and at first he treated his two lots
of material as Maskell’s species. He defines it as a new species on the structure
of the pygidium, in the shape of the trullae and lobules of the adult female, which
he figures. The specimens were collected upon undetermined species of Casuarina
from Biniguy, New South Wales, and Chinchilla, Queensland, by Mr. Gerald F.
Hill.

POLIASPIS CASUARINAE Lidgett.

This species was described by Mr. Lidgett upon the branchlets of Casuarina
suberosa growing at Myrniong, Victoria. I have given the author’s description of
this species in my Descriptive Catalogue (pt. 1, 1915, p. 49). As the type is, I
understand, missing, its determination without fresh specimens from the exact
locality will be difficult.

FIORINIA CASUARINAE Maskell.

This species comes from near Perth, W. Australia, upon the branchlets of an
undetermined species of Casuarina. The puparium of the female white, of the
typical elongate form. About 1/12 inch in length. Adult female brown, very
small. Male puparium like that of female, but with only one pellicle.

ALECANOPSIS (LECANOPSIS) CASUARINAE Mask.

The adult female is a semiglobular reddish-brown coccid with the outer
margins black, fitting close against the wood, with the under surface convex,
forming a box-like cavity. Diameter 3 inch. ;

The type specimens were taken out of the deserted burrow of a wood moth in
the centre of a stem of a sheoak (Casuarina sp.) in which there was a colony of
ants. Hab.—Myrniong, Victoria.

This coccid is now placed in Cockerell’s genus Alecanopsis, the members of
which are lecanids which have adopted the habit of living in cavities in tree
trunks, or on roots, and are usually associated with ants, which also form nests in
the same bores.

Maskell’s Lecanopsis filicum is the type of the genus; it was collected on the
Kurrajong Hills, N. S. Wales, upon the roots of a fern, Woodia aspera Green
(Bull. Entom. Research, ii, 1924). Three more species from Australia: A. tenuis
from Victoria in the stems of Banksia integrifolia, and Green identified specimens
which I collected in the stem of Grevillea robusta at Bulga, N. S. Wales, as the
species. Both of these were attended by ants. A. merus, from Townsville, Queens-

BY W. W. FROGGATT. 365

land, was found in an ants’ nest (Cremastogaster australis) in a tree trunk.
A. grandis was described from specimens on a fern root, in the British Museum
collections from Bundarra, N. S. Wales. An attached note recorded that it was
taken out of a nest of Campinotus intrepidus. The three species which I described
in These ProckEepINGs, 1925, will also come into this genus.

RHIZOCOCCUS CASUARINAE Maskell.
Trans. N. Zealand Inst., xxv, 1892, p. 230, Pl. v, fig. 7.—The Entomologist, xxvii,
1894, p. 46.
This coccid is recorded from Myrniong, Victoria, upon Casuarina suberosa,
and from Cheltenham, Victoria, upon Casuarina distyla.

RHIZOCOCCUS LECANIOIDES Green.
Bull. HEntom. Research, vi, pt. 1, 1915, p. 47, fig. 4.
This species comes from Sandringham, Victoria, upon Casuarina distyla.

Rurzococcus MANcUS Maskell.
Trans. N. Zealand Inst., xxix, 1897, p. 316.
This is the common species about the Sydney coast upon several species of
sheoaks, Casuarina suberosa, C. distyla and C. rigida. It is figured in my Cat.
Coccidae, pt. ii, p. 66, fig. 48.

RHIzZOCOCcUS PUSTULATUS Maskell.
Trans. N. Zealand Inst., xxv, 1892, p. 231, Pl. xv, figs. 8, 9.
This is another species recorded from Myrniong, Victoria, on an undetermined
species of Casuarina.

RHIZOCOCCUS TRIPARTITUS Fuller.
Journ. W. Aust. Bur. Agric., iv, 1897, p. 1345.—Trans. Ent. Soc. London, 1899,
p. 448.
This was described from Western Australia on an undetermined species of
Casuarina.

GOSSYPARIA CASUARINAE Maskell.
Collected in the vicinity of Sydney, upon the branchlets of an undetermined
species of Casuarina.
The adult female coccid varies from light to dark brown in colour, is about 1/12
inch in length, and rests upon a pad of woolly matter with the dorsal surface
exposed.

ERIOCOCCUS CONSTRICTUS, nN. sp. Text-figs. 4-6.

These coccids are solitary, the sacs being scattered over the branchlets of
Casuarina lepidophloia, Huston, Murray River, N.S.W.

Adult °. Sac light reddish-brown, surface smooth, stout, and closely felted;
general form elongate-oval, with the apical portion slightly contracted, the anal
orifice large, circular. The sac attached to the branch along the ventral surface
with the anal tip turning upward. Length 4 mm., height 2 mm.

Adult 9 coccid dark brown, giving amber tints when boiled in potash. Dorsal
surface rounded, ventral surface flattened, extremities rounded. Length 34 mm.
Antennae 7-jointed, rather short, joint 1 broad, 2 rounded at apex, 3 larger, 4
shortest, 5 slender, longer, 6 slender, shorter, 7 about the same length with
several fine hairs at the extremity. Legs well developed, femora short and broad,
tibia broad, tarsus long, upper edge bearing long hairs, under-surface with shorter

Co)

366

THE COCCIDAE OF THE CASUARINAS,

EA King

Text-figs. 1-3.—Hriococcus fossilis. 1, group of sacs on branchlets;
2, showing form of sac; 3, ventral surface of adult female.

Text-figs. 4-6.—Hriococcus constrictus. 4, sacs on branchlet; 5, showing
form of sac; 6, ventral surface of adult female.

Text-figs. 7-9.—Hriococcus rugosus. 7, sacs on branchlet; 8, showing
structure of sac; 9, ventral surface of adult female.

Text-fig. 10.—Sphaerococcus ethelae. Ventral surface adult female.

Text-figs. 11-12.—Casuarinaloma leai. 11, showing section of gall;
12, upper surface of gall.

Text-figs. 18-14.—Sphaerococcus ethelae. 13, larva; 14, showing aborted
tissue of galled wood.

BY W. W. FROGGATT. 367

hairs, claw small. Derm clothed with fine pores, some irregularly rounded and
fine spines. Anal lobes produced into a pair of large angular plates, slightly
arcuate on the inner margins, with six hairs, and several spines on, the inner
edges.

ERriococcus CYPRAEAEFORMIS Fuller.
Journ. W. Aust. Bur. Agric., 4, 1897, p. 1845.—Trans. Ent. Soc. London, 1899,
p. 440.
This curious Hriococcus comes from Western Australia, exact localit'y not
given, upon Casuarina sp. The figure in my Cat. Coccidae, part ii, p. 79, gives
a good idea of the distinct shell-like form.

Eriococcus ELEGANS Fuller.
Journ. W. Aust. Bur. Agric., 4, 1897, p. 13845.—Trans. Ent. Soc. London, 1899,
p. 440, Pl. xii, fig. 4.
This species comes from near Perth, W. Australia, upon Casuarina humilis.

ERIOCOCCUS FOSSILIS, n. Sp. Text-figs. 1-3.

The specimens were collected by Mr. G. F. Hill upon the branchlets of the
river oak, Casuarina OCunninghamiana, Paddy’s River, Canberra.

The adult ¢ sacs cluster together towards the tips of the branches; they are
deep biscuit-brown, with the surface very smooth, oval in form, the anal opening
very large and raised above the lower margin of the sac. Length 3 to 4 mm.

Larva pink, broadly oval with long six-jointed antennae, legs long and slender,
bearing a fine hair and two clubbed digitules; no marginal spines on the cephalic
or thoracic segments, but the abdominal ones with a fringe of eleven spines
increasing in length to the anal lobe. The anal segment produced into two
projecting tubercles each bearing one long and two shorter bristles between them,
and a fringe of about ten very fine hairs.

Adult 2 coccid brown, general form oval. Length 34 mm., width 2 mm.
Antennae slender, 6-jointed, 1 small, 2 short, broad, 3 twice the length of 2, 4 and
5 short, irregularly rounded, 6 longer than 5, arcuate and somewhat irregular in
form. Legs moderate, tibia short, tarsus long, terminating in a small sharp,
curved claw. Derm lightly clothed with fine spines, with larger spines on the
outer margins of the abdominal segments. Segments 4—7 with the upper margins
banded with brown chitin, the last three bands angulated. Pygidium large,
rounded behind, chitinous, closely shagreened, lobes spade-shaped, serrate on the
inner edges, with a fringe of fine hairs between them, outer margins bearing two
spines.

ERIOCOCCUS RUGOSUS, n. sp. Text-figs. 7-9.

Collected upon the branchlets of Casuarina Luehmanni, Murray River, near
Euston, N. S. Wales, scattered over the branchlets usually in groups of two or
three.

©. Puparium dull reddish-brown, short, and broad in proportion, composed of
closely felted filaments with the surface slightly roughened, anal extremity
truncate, opening large. Length 3 mm.

Adult 2 brown, giving off purple tint when boiled in potash. The dorsal
surface broadly rounded. Length 24 mm. Derm thickly covered with fine short
spines. Antennae slender, composed of 7 segments, tapering to the tips, 1 short
and broad, 2 and 8 cylindrical, longer than the first, 5 and 6 bead-shaped, of about
equal length, 7 slightly longer, contracted at the tip, clothed with a few fine

368 THE COCCIDAE OF THE CASUARINAS,

hairs. Legs slender, 1st and 2nd pair with the tibia slender, hind pair stouter,
tarsus with small tarsal claw. Pygidium forming a pair of projecting triangular
ochreous plates with six hairs between them.

PSEUDORIPERSIA BREVIPES, N. sp. Text-figs. 26-28.

This fine species, the second of the genus, comes from Euston, Murray River,
N. S. Wales, where I collected it upon the branchlets of Casuarina lepidophloia.

The saes are clustered together in groups of from 5 to 30 upon the smaller
branches. They are composed of grey-coloured felted secretion, of much softer
material than those of P. turgipes, forming a cup-shaped sac in which the 9 coccid
rests, covered with a thin white pellicle, above which is a conical cap of similar
material to that of which the sac is composed; it fits within the cup, projecting
well above the rim, and is not attached to the basal sac. Height about 3 mm.

Adult 2 coccid 2 mm. in diameter, subglobular, dorsal surface flattened but
showing a slight marginal ridge. Colour dark brown, when boiled in potash
giving off a deep green colour. Antennae 6-jointed, 1 and 2 short, 3 longest, 4
and 5 very short, 6 nearly as long as 3. The antennae shorter and smaller than
those of P. turgipes. Legs similar in general form but shorter. The derm with
scattered pores, and irregular bands of spines and pores forming a band round
the cephalic, thoracic and abdominal segments; with irregular bands of spines
across the lower margin of the abdominal segments. The anal segment, with a
pair of pear-shaped lobes between which is the anal ring, which is fringed with
stout hairs; a few long spines on either side.

PSEUDORIPERSIA TURGIPES Maskell. Text-figs. 23-25.

This coccid has a wide range and infests the branchlets of a number of
different species of Casuarina, sometimes singly, but often in groups. The
presence of the coccid often causes the branchlet to curve right round. In the
vicinity of Sydney they infest Casuarina suberosa; in western New South Wales
and southern Queensland it is common on the Belah (C. Cambagei).

Sac almost globular, 3 mm. in diameter, composed of leathery white secretion
slightly roughened on the surface. The base attached to the branchlet, and the
apex with a large circular opening in the sac, beneath which the coccid rests,
with a convex pad of similar white secretion not attached to the sac, protecting
the enclosed coccid and level with the top of the sac.

Morrison gives a detailed account of the adult coccid and larva on page 51
of his “Maskell’s Genera of Coccidae’’, and figures them (fig. 14).

PSEUDOCOCCUS CASUARINAE Maskell.
This species was described from specimens found upon the stems of an undeter-
mined species of Casuarina at Myrniong, Victoria.
The adult female is enveloped in an irregular mass of white cottony substance.
Yellowish-brown, dorsal surface convex. Length 2 inch.

CASUARINALOMA, N.g.

The female coccids forming and living in galls. The male coccids developing
within the gall chamber, enclosed in a fine silken sac, usually three or four in
number.

Adult 2 coccid with well developed legs.

CASUARINALOMA (SPHAEROCOCCUS) LEAI Fuller. Text-figs. 11, 12, 20.
Fuller described the type from galls on an undetermined species of Casuarina
collected near Perth, W. Australia. Mr. J. Macqueen sent me specimens from

BY W. W. FROGGATT

PALL AMUOVITITI RTA

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SAPNA S PAAR’

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Text-figs. 15-17.—Cylindrococcus spiniferous.

Text-figs. 18-19.—Frenchia caswarinae. 18, adult female; 19, anal
segment.

Text-fig. 20.—Casuarinaloma leai. Ventral surface, adult female.

Text-figs. 21-22.—Gymnaspis serratus. 21, dorsal surface, adult female;
22, ventral surface.

Text-figs. 23-25.—Pseudoripersia turgipes. 23, cluster of sacs; 24, single
sac, viewed from above; 25, ventral surface, adult female.

Text-figs. 26-28.—Pseudoripersia brevipes. 26, group of sacs; 27, sacs
viewed from above; 28, ventral surface, adult female.

369

370 THE COCCIDAE OF THE CASUARINAS,

Millmerran, Queensland, from Casuarina sp. I found them plentiful upon
Casuarina Cambagei at Trangie, N. S. Wales, and upon the branchlets of
C. Luehmanni near Euston, Murray River, N. S. Wales.

The galls are in clusters of three or four, or singly upon the tips of the
branchlets; they measure 6 mm. in diameter, flattened on the summit with the
sides deeply cleft into from 12 to 10 segments, forming a circular solid but thin-
walled gall which somewhat resembles the seed capsule of a marsh-mallow.

&. Coccids developed inside the gall-chamber; in matured galls there are
often three or four white oval silken puparia open at the apex. Males red, wings
white, head broad, the antennae composed of small irregular segments, fringed
with hairs; body short, segments slender, terminal one lance-shaped.

@. Coccid broadly rounded, convex, rounded on the ‘dorsal surface. Diameter
14 mm. Colour red. Antennae composed of six small somewhat irregular joints.
Legs well developed, femur large, rounded, tibia short, broad; tarsus long,
terminating in a‘fine curved point. Derm thickly covered with stout finely-pointed
spines, no distinct pores, but the whole surface finely shagreened. Anal lobes
very small, inconspicuous, bearing three hairs, the central one longest, the anal
ring apparently between them in a slight depression.

Larva: I have never seen the larva, but Fuller says, “Larva elongate fringed
with spines. Anal tubercles bearing setae and spines. Antennae 6-jointed. Legs
thick, tarsus slightly longer than tibia. Upper and lower digitules knobbed.
Colour crimson.”

A bunch of the galls is figured in my Descriptive Catalogue of Australian
Coccidae, part ii, 1921.

Fuller placed this species in Maskell’s genus Sphaerococcus, but this genus
is defined “legs absent or atrophied”, so that the possession of three pairs of well
developed legs places this species outside this genus.

FRENCHIA CASUARINAE Maskell. Pl. xxix, fig. 4; Text-figs. 18, 19.

The type specimens came from Victoria, on the stems of Casuarina
quadrivalvis. My specimens were collected by Mr. C. French, Jr., on the same
species of sheoak growing near Geelong, Victoria. I have also found this species
in New South Wales on an undetermined species of sheoak, but it is a rare gall
in this State. It is found in Tasmania on C. quadrivalvis. These galls are
gregarious; they are usually in numbers on each infested branch, often so close
to each other that the basal swollen wood tissue or false galls are in contact.
This swollen tissue with the covering bark surrounds the true gall in which the
coccid is imprisoned.

This gall is a rounded, smooth, dark brown, hard, wooden tube, projecting
above the swollen tissue about 10 mm. The apex is nipple-shaped, ringed, and
light reddish-brown; the apex closed with a truncate circular plug, which protects
the coccid until the larvae are ready to emerge. The gall is about 4 mm. in
diameter in the centre, but at the hidden base much enlarged, fitting against the
sapwood; it is 8 mm. in diameter; the combined length of basal stem and erect
tube is about 15 mm. When the tubular gall is cut out and detached there is a
circular scar in the sap-wood beneath. Here, fitting into the base of the tube
between the scar and the cephalic portion of the adult female, is a rounded,
yellow, waxy object, with a curved horn-like process in the centre. Maskell
called it the indusium, or third gall. Morrison, in his examination of Maskell’s
material, said that Maskell was in error; he says: “The second stage as described

BY W. W. FROGGATT. 371

by Maskell is in reality a first stage larva about to moult.’ I have had a great
number of fresh galls through my hands, and in every one containing a live
adult female, this second stage larva is present.

Adult 2 is reddish-yellow with a little white floury meal scattered over it ou
the under-surface, but more thickly coated on the upper surface. The cephalic

E.AKing

Text-figs. 29-36.—Cylindrococcus amplior. 29, showing section of gall;
30, group of galls; 31, ventral surface, adult female; 32, dorsal surface, adult
female; 33, antennae, adult female; 34, anal ring; 35, ventral surface of
larva; 36, dorsal surface of larva.

372 THE COCCIDAE OF THE CASUARINAS,

portion coalesced with the thoracic segments forming a thickened rounded disc,
the abdominal segments contracted into a slender tapering tubular tail. Diameter
of disc 6 mm., thickness 1 mm., abdominal tail from disc to tip 6 mm. When
alive it might be likened to a slender-stalked mushroom. Morrison, in his definition
of the genus Frenchia, places it in the subfamily Dactylopinae; he figures the
larvae and details of the adult coccid, and concludes with a diagnosis of the
genus.

FRENCHIA SEMIoccULTA Maskell. Plate xxix, fig. 5.

The galls upon which Maskell formed this species were collected by the writer
upon the branchlets of Casuarina suberosa at two localities near Sydney. In
this species the galls form a double swelling on the branchlet like two lips, with
a cleft between them, at the base of which is a cavity containing the coccid. The
specimens figured are immature galls. I collected a somewhat similar gall on
the branchlets of Casuarina Luehmanni near Euston, on the Murray River, N.S.W.

The adult 9 coccid resembles the preceding species in general form and
colour, but is much smaller and the abdominal segments are not so elongated. In
Morrison’s paper, “The Maskell Species of Scale Insects of the Subfamily
Asterolecaniinae”’, this species is described at length on pages 22-24, and the
details are given on Plates 14 and 15.

SPHAEROCOCCUS ETHELAE Fuller. Text-figs. 10, 13, 14.

The @ coccids are gregarious, their presence causing scars upon the branches
of several species of Casuarina; they can hardly be called gall-makers. Fuller
described the type from the branches of a Casuarina, Swan River, W.A.

Collecting on the banks of the Murray River, near Euston, N. S. Wales, I
found many of the stunted sheoak, Casuarina Luehmanni, with the branches
infested with abandoned galls of Frenchia semiocculta, many branchlets having
snapped off at the double-lipped gall. These damaged tips were aborted in irregular
scars covered with white flocculent matter looking like woolly aphis. These
woolly filaments covered several coccids further protected with thin silvery-
white plates, the last folded over the coccid.

©. Coccid 13 mm. in length, chocolate-brown, when treated with potash giving
out a bright green tint. Derm thickly covered with small circular pores. Antennae
small, indistinct. Legs rudimentary, four in number, represented by two pairs
of raised irregularly rounded coriaceous plates, from which project short tubular
processes (tibiae?) with a fine prong on either side. Anal plate chitinous, basal
portion finely granulated. Anal lobes short, broad, and projecting slightly.

Larva found in cavity with adult female of very distinctive form as figured.
Further investigations into the structure of this coccid may place it in a new
genus. All the coccids in the galls were dead and dry.

CyYLInpRococcUS AMPLIOR Maskell. Pl. xxix, fig. 1; Text-figs. 29-36.

The type specimens were sent to Maskell from near Adelaide, S. Australia, on
the branchlets of Casuarina quadrivalvis. I have specimens from Glen Osmond,
S. Australia (J. Davidson), on C. stricta; Victoria (C. French, Jr.), on
C. quadrivalvis; Millmerran, Southern Queensland (J. Macqueen), on (©. torulosa;
Gilgandra, N.S.W. (G. A. Withers), on stems of Casuarina sp.

The drawings were made from living unmounted specimens as well as
mounted specimens. In general form and structure, and enfolding of the larvae,
the adult female resembles C. casuarinae. It is somewhat larger and broader,
measuring 6 mm. in length and 4 mm. in diameter. Of the same reddish-brown

BY W. W. FROGGATT. 373

tint, with the colour obscured by white floury secretion. The antennae cone-
shaped, composed of 4 segments, with single spines on the sides of the apical
ones. The dorsal surface showing ten segmental divisions, the apical one broadly
oval, chitinous, brownish-yellow, thickly covered with fine spines, with the margin
fringed with longer stouter ones. A few larger spines across the other abdominal
segments, but they are not fringed with fine yellow spines as in (C. casuarinae.
The ventral surface very similar. Anterior legs short and aborted, with the
prolonged lobes of the thoracic segments ringed and broadest at the base; the
lower pair not projecting. The anal ring distinct, with two stout spines. The
larvae do not appear to differ from those of C. casuarinae. The galls are apparently
also formed from a branchlet bud, with the leaves forming a cup of elongated
bracts like an acorn, and not attached to the sides of the central gall. This is
reddish-brown, but the surface is covered with greyish down. Smooth, broad, and
round at the base, it has the sides ribbed, lightly at the base, but more distinctly
channelled to the pointed apex. They vary from 20 to 25 mm. in length, and
from 5 to 12 mm. in diameter. The gall chamber is slender and oval, but the
walls are solid at the base, and thickened on the sides to the extremity.

CYLINDROCOCCUS CASUARINAE Maskell. Pl. xxix, fig. 2.

The type specimens were collected near Melbourne, Victoria, upon the
branches of Casuarina quadrivalvis. I have examined a number of living and
mounted specimens taken from fresh galls on the same species of sheoak collected
near the same locality by C. French, Jr., this year.

I have also recently taken this gall upon the foliage of Casuarina stricta
growing on a headland near Newport, N. S. Wales.

Morrison, in his paper, “A redescription of the type species of the genera of
Coccidae based on species originally described by Maskell’, considers this species
the type of the genus Cylindrococcus. In describing it, he figures the larva and
adult female with details of the external appendages. His descriptions are based
upon material boiled in potassium hydroxide; I would give some notes on the
general appearance of the living coccids when removed from the galls, with an
account of the origin and structure of the galls.

Adult female dark reddish-brown, but smothered in white floury secretion,
which is easily rubbed off. They vary in size from 43 to 5 mm. in length, and
2 mm. in diameter, and rest loosely on the lower half of the gall chamber, with
the ventral cavity packed with hundreds of minute dark red larvae, which crawl
out all over the paper when the coccid is laid on her back.

The general form is cylindrical, rounded at the extremities, with the 4-
jointed conical pointed antennae projecting above the apex of the cephalic segment,
but not as much as in C. amplior. The dorsal surface is rounded, showing eleven
segmental divisions; the cephalic rounded, 1st thoracic largest, 2nd and 3rd about
the same size, with the first six abdominal segments narrow and of uniform width;
the 7th forming a smooth circular conical plate, coriaceous, when examined under
the lens seen to be covered with very fine spines which fringe the outer margin and
cross the six abdominal segments; it curves round to the ventral side, and with
the outer margins of the six abdominal segments folds over, forming the well
defined pouch or cavity previously noted, in which the larvae assemble and
remain for some time before emergence. Viewed when resting on its back, it
might be likened to a little rubber doll, the antennae projecting like ears, the
anterior legs, spiracles and impressed rostrum forming the face. The ist thoracic
segment is swollen out on either side into a nipple-like process in the immature

D

374 THE COCCIDAE OF THE CASUARINAS.

female, but in the adult is flattened, rounded at the extremity and folded over
the side of the body, the 3rd thoracic segment with a similar projecting process
but much smaller. These extraordinary appendages may represent legs, as
Morrison says, but I am inclined to think the adult females have only a pair
of forelegs.

Galls: These are sessile, springing out from the ringed ridges on the smaller
stems, and appear to be developed from a branchlet bud, and the basal bracts or
scales on the lower third of the gall represent the rudimentary leaves which
encircle the normal branchlet.

When there are several galls on the same node, the pointed bracts form an
expanded fringe round the base, but in solitary galls they fit more closely round
the surface of the gall. The galls are faintly-ribbed thin tubes, closed at the
extremities until they are quite mature.

CYLINDROCOCCUS SPINIFEROUS Maskell. Pl. xxix, fig. 3; Text-figs. 15-17.

This is the most abundant gall along the foreshores of Sydney Harbour, in
the early summer, covering the branchlets of Casuarina suberosa with its bracteate
seed-cone-like galls. Though their structure is identical, they vary very much in
form during their development. The pointed bracts apparently produced from
the rudimentary leaves being thin, and fitting over each other like a small pine
cone with the pointed tip of the thin central tube enclosing the female coccid
projecting at the apex. In the old mature galls, the bracts are thickened,
coalesced at the base, forming a squat gall, with the apex depressed-convex and
perforated in the centre.

The type specimens came from Victoria. Mr. Charles French collected them
upon Casuarina quadrivalvis near Melbourne. It is recorded upon the same species
from South Australia and New South Wales.

The living larvae are bright red, and cluster together in the cavity on the
ventral surface of the mother. They are shield-shaped, but more elongate and
rounded at the apex than the other two species, with similar 4-jointed antennae
and legs, but the fringe of fine spines along the sides is not so prominent, though
the outer long spines on the anal lobes are slightly longer.

Adult 9. Length 6 mm., diameter 4 mm. Red, with the tip of the abdomen
black; the ventral thoracic segments sometimes variegated with black spots.
General form cylindrical, apex truncate, broadest, deeply wrinkled and pitted.
Dorsal surface smooth and rounded, showing five large and four narrow red
segmental divisions, with one fine black one round the black circular convex
tip, which is clothed with fine spines. The ventral surface with the cephalic
region much wrinkled and contracted; the thoracic segments smooth, swollen on
the sides, but without the nipple-like projections of the immature females.
Impressed in the centre with a smooth angular area above the first abdominal
segment. Abdominal segments depressed and flattened on the central area, but
the outer margins raised, consisting of six narrow segments, with the circular
convex tip turning inwards forming an abdominal cavity in which the larvae are
massed together.

EXPLANATION OF PLATE XXIX.
1. Galls of Cylindrococcus amplior.
2. Galls of Cylindrococcus casuarinae.
3. Different forms of galls of Cylindrococcus spiniferous.—Sessile, stalked and upon
seed-cone.
4. Galls of Frenchia casuarinae.
5. Galls of Frenchia semiocculta.

Proc. Linn. Soc. N.S.W., 1938. PLATE XXIX.

Galls of (1) Cylindrococcus amplior, (2) C. casuarinae, (3) C. spiniferous,
(4) Frenchia casuarinae, and (5) F. semiocculta.

AN INVESTIGATION OF THE SOOTY MOULDS OF NEW SOUTH WALES. I.
HISTORICAL AND INTRODUCTORY ACCOUNT.
By Livranw Fraser, M.Sce., Linnean Macleay Fellow of the Society in Botany.
[Read 25th October, 1933.

1. Introduction.

2. Historical Review: A. Systematic (i, Capnodiaceae; ii, Atichiaceae; iii, Fungi
Imperfecti). B. Physiological.

3. Sooty Moulds of N.S.W.: A. Fungi Recorded (i, Capnodiaceae; ii, Atichiaceae ;
iii, Fungi Imperfecti). B. Development and Distribution.

4. The Problem of Polymorphism: A. Historical Review. B. Cultural Evidence.
5. Conclusions.

6. Scale Insects Concerned.

7. Summary.

Introduction.

The term “Sooty Mould” is used to describe the dark brown or black felt
of fungal mycelium often found on the leaves and branches of plants attacked by
scale insects or aphis. The corresponding French and German terms are
“Fumagine”’ and “Russtaupilze”.

The fungi composing the mould are saprophytes which utilize the honey-dew
secreted by scale insects and aphis. They therefore do no direct harm to the
plant on which they occur, but may damage it indirectly by interfering with
photosynthesis.

Sooty mould fungi are always strictly superficial and do not develop haustoria.
The mould only adheres to the host leaf by virtue of the mucilaginous nature of
the cell walls of the fungi composing it, and therefore flakes off readily when dry.

As well as being common on trees attacked by scale insects, sooty moulds
have been recorded on the extrafloral nectaries of Hibiscus in Java by Boedijn
(1931), and of cotton in India by Sawhney (1927).

It was originally thought that any particular sooty mould consisted of a
single fungus in the same way that a particular kind of plant disease was due
to a single pathogenic fungus. It is now known that in practically all cases
sooty moulds consist of a number of fungi which live together forming a colony,
and fruit concurrently. These different fungi have in common a dark-coloured
mycelium, the cell walls of which have a tendency to become mucilaginous when
wet.

The fungi which have been found to be of importance in the formation of
sooty moulds can be classed into three groups: 1, Those which belong to the
Capnodiaceae. 2, Those which belong to the Atichiaceae. 3, Those which belong
to the Fungi Imperfecti.

To the Capnodiaceae belong the characteristic members of the sooty mould
flora. In tropical countries where sooty moulds are particularly abundant it is
not uncommon to find the perfect or imperfect fruits of as many as seven
members of the Capnodiaceae in a single mould (Mendoza, in Stevens, 1925). It

376 THE SOOTY MOULDS OF NEW SOUTH WALES. i,

is a striking fact that the most abundant and conspicuous fungi of sooty moulds
should thus form a natural group systematically. They grow exclusively on the
honey-dew of scale insects.

The members of the family Atichiaceae are aberrant saprophytic ascomycetes
which grow on the leaves of trees, principally in the tropics. The most wide-
spread species is Atichia glomerulosa, which is common in sooty moulds through-
out Europe.

Those members of the Fungi Imperfecti which are found in sooty moulds
are usually common saprophytic species such as Cladosporium, Dematium and
Penicilium spp.

Sooty moulds are found throughout the world, but they differ widely in
character in different localities. The maximum development is reached in wet
forest areas, especially in tropical countries where rainfall is heavy and the
atmosphere humid. Members of the Capnodiaceae and Atichiaceae are most
numerous in the tropics. They are less important in cold temperate regions.
Here the Fungi Imperfecti are plentiful and may be present in the sooty moulds
to the exclusion of the Capnodiaceae and Atichiaceae.

A moist atmosphere is particularly favourable for the Hevelonment of sooty
moulds. Zopf (1879) and Thuemen (i890) noted their frequent appearance on
the leaves of green-house plants. Thuemen recorded that in Hurope their principal
development takes place in the months July to November, especially in the late
autumn, and stated that the amount of mould increased proportionately with the
atmospheric humidity. Neger (1918) dealt in great detail with the necessity for
a moist atmosphere for the development of sooty moulds. Mendoza (1932) pointed
out that in the Philippine Islands where sooty moulds are very abundant, the
greatest development takes place in the rainy seasons.

HISTORICAL REVIEW.
A. Systematic.

Much confusion of nomenclature exists in the early descriptions and discus-
sions of sooty mould forming fungi. This is due to several causes, the chief of
which has already been noted. This was the tendency to regard any particular
development of sooty mould as being formed of a single polymorphic fungus which
produced a great variety of fructifications. A few workers only considered
that each type of fructification found in a Sooty) mould was produced by a
different fungus.

Another cause for confusion of nomenclature was the view, at first widely
accepted, that the species of fungus causing a sooty mould differed with the host
plant, as with parasitic fungi. Consequently identical fungi were often given
different names if they were found on different host plants.

In many cases, too, descriptions of type species and genera were inadequate,
and consequently were variously interpreted by later workers. The result was
that a very heterogeneous group of fungi was in many cases attributed to the
one genus.

A brief summary is given below of some of the most important changes in
nomenclature of the more widespread and better known mould-forming species of
fungi.

(i) Capnodiaceae.
Amongst the earliest sooty mould genera to be described were Antennaria Link
in 1809, Apiosporium Kunze in 1817, Fumago Pers. in 1822, and Scorias Fries in

BY LILIAN FRASER. 377

1825. During the following few decades many species were ascribed to these
genera, most of them on inadequate grounds.

Amongst the most important of the species described were Fumago citri
Turp., the “citrus mould’, and Fuwmago salicina Pers., the “willow mould’, and
Fumago vagans Pers., which was stated to be widespread.

In 1849 Montagne described the genus Capnodium to include Fumago salicina
‘and F. citri on the ground that the type species of Fumago was based on an
imperfect fructification. Capnodium was described as having club-shaped
perithecia, and 4-septate, yellowish-brown ascospores.

Tulasne (1863) retained the older nomenclature. In describing Fumago
salicina Pers., he united under this name some 14 fungi which had been described
as different species. Some of the species included by him were Fumago vagains
(in part) and other species of Fumago, Cladosporium spp. and Fumago citri (i.¢.,
Capnodium citri of Montagne). He also considered that Antennaria elaiophila
Mont. (which is the same as TJorula oleae Castaigne) and Antennaria semi-ovata
B. & Br. are probably identical with Fumago salicina.

Tulasne placed the Fumagines in the Sphaeriei.

Farlow (1876) described the sooty mould of oranges in California as belonging
to the species Capnodium citri Mont. He expressed the opinion that Capnodium
citri and the fungi described as Fumago citri Pers., Antennaria elaiophila Mont.
and Fumago salicina were all identical, thus confusing two very distinctive
fungi, Capnodium citri and C. salicinum.

Zopf (1878) described in detail the development in culture solutions of a
sooty mould fungus common at that time in the glass houses in Berlin, growing
saprophytically on the secretions of aphis and scale insects. This fungus he
classified as Fumago, but since he found no perfect fruit and none developed in
his cultures, he was unable to assign it to any definite species. Zopf found that
this species was extremely polymorphic. In pure culture it produced a large
number of different kinds of fruiting structures showing all gradations from
free conidiophores to closed pycnidia. This work appeared to confirm the general
opinion that all sooty moulds are polymorphic.

Penzig (1882) described the sooty moulds of citrus, which had been studied
intensively because of the economic importance of the host plants, as belonging
to the genus WMeliola, as Meliola citri. This classification was also used by
Saccardo who considered that there were three species of Meliola which formed
sooty moulds on citrus, Meliola citri, M. camelliae and M. Penzigi. Penzig
considered that the two latter were probably identical.

Gaillard (1892), in a monograph of the genus Meliola, excluded from this
genus the fungi responsible for the sooty moulds of citrus.

Weber (1896) retained the older classification of Saccardo, referring the
sooty moulds of citrus in Florida to the species Meliola camelliae.

Lindau, in “Die Naturlichen Pflanzenfamilien” (1897), made further changes
in nomenclature. He included the species previously known as Capnodium and
Fumago in the genus Apiosporium Kunze, except for five species of Fumago
retained in the Fungi Imperfecti near Alternaria. The genus Antennaria Link
was retained and in it was included Limacinia, a genus described by Neger in
1896, as forming a sooty mould on the leaves of trees in Juan Fernandez.
Apiosporium, Antennaria and the sooty mould Scorias spongiosa were included
by Lindau in the Perisporiaceae with Meliola and Perisporium.

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THE SOOTY MOULDS OF NEW SOUTH WALES. i,

Another change in nomenclature was later made by Saccardo, who placed the
sooty moulds of citrus in the genus Limacinia Neger, since the perithecia produced
by them were sessile. He limited Capnodium to those species with stalked or
vertically extended cylindrical or club-shaped perithecia, of which the type is
C. salicinum (see von Hoehnel, 1909b, and Arnaud, 1911). In contrast to Lindau,
Saccardo considered that Antennaria is the unripe fruit of Capnodium, and that
Capnodium is not identical with Apiosporium. He placed the sooty mould genera,
including Limacinia, Scorias, Capnodium, etc., in the Capnodeii, a section of the
Perisporiaceae.

Lindau’s scheme of classification was used by Sorauer (1908) in his handbook
of plant diseases, and is retained in the later editions.

Clements (1909) largely followed the arrangement of Saccardo, but placed
Limacinia as a synonym of Meliola, evidently ignoring its saprophytic nature.

Von Hoehnel (1909a) pointed out that the genus Apiosporium Kunze was
invalid, since the type specimen is a sclerotium. He reviewed the genus in
detail and reclassified the species referred to it into a number of other genera. He
also reviewed the genus Antennaria at some length. The description of the type
species of Antennaria, A. ericophila Link, is said by von Hoehnel (19090) to be
inadequate. According to him most of the species subsequently placed in this
genus were incompletely known fungi of doubtful position. The type species
was redescribed by Neger, and Neger’s fungus has been shown by von Hoehnel
to be the imperfect stage of Coleroa Straussii (Sacc. & R.) v. H., a parasitic species
of the Sphaeriales. According to von Hoehnel, therefore, Antennaria Link becomes
Coleroa Rab. In any case the generic name Antennaria is invalid, since it had
been applied previously to a genus of the Compositae by Gaertner, and was super-
ceded by the name Antennularia of Reichenbach in 1828 (see von Hoehnel, 1909a).

Since then the name Antennularia has been used by von Hoehnel to designate
the pear-shaped pycnidia of Coleroa and the sooty mould genus Limacinula
(1909b). It has been used by Woronichin (1926) and Boedijn (1931) for the
pycnidial fruits of various members of the Capnodiaceae.

In describing a new sooty mould, Limaciniula samoensis, von Hoehnel (1909b)
reviewed the genera and species of the Capnodeii as described by Saccardo. He
considered that the sooty mould fungi which constitute this section form a very
natural group. He maintained, however, that the genus Limacinula should not be
placed in the Capnodeii, and suggested the formation of a new family, the
Naetrocymbeii, to include it, this family to be placed in the Sphaeriales. The
Naetrocymbeii were said to parallel the Capnodeii in having similar mycelium
and conidia, but to differ in the formation of the perithecium which is never
cartilaginous and tough as in the Capnodeii, but is soft and membranous with a
well developed ostiole. There were two genera in this new family: Naetrocymbe
Korb. and Limacinula Sacc., the type being Limacinula javanica (Zimm.) Sacc.

In 1910 von Hoehnel raised the subsection of Saccardo to the position of a
family, the Capnodiaceae, and gave a scheme of classification.

A very different attitude was adopted by Arnaud (1910a, 19106, 1911, 1912).
He did not consider that the genera placed by von Hoehnel in the Capnodiaceae
formed a natural group, but maintained that they are members of the Sphaeriaceae.
The individual species of sooty moulds he re-classified into already known genera
of the Sphaeriaceae, Pleosphaeria and Teichospora. Limacinula, Capnodium and
Teichosporina were taken as subsections of the genus Teichospora. The genera
Limacinia and Scorias were retained and transferred to the Sphaeriaceae.

BY LILIAN FRASER. 379

Arnaud reviewed in considerable detail the sooty moulds described from citrus
plants, and dealt very adequately with the confusion of nomenclature existing
in this group. He gave a full.list of synonyms. The result of his review was
that he placed the fungi previously referred to as Limacinia Penzigi, L. citri and
L. camelliae by Saccardo (i.e., Fumago citri Turp., Capnodium citri Mont., and
Meliola citri, M. camelliae and M. Penzigi Sace.) in the genus Pleosphaeria. He
considered that all are identical and renamed the species Pleosphaeria citri Arn.
Some of the imperfect fruit attributed by older authorities to this fungus were
considered by Arnaud to belong to a newly described sooty mould Teichospora
meridionale Arn.

Arnaud’s scheme of classification was not accepted by contemporary German
mycologists, and in 1916 Theissen put forward a new classification, retaining the
family Capnodiaceae as a division of the Perisporiales. He also expressed doubts
as to the validity of the family Naetrocymbeii as described by von Hoehnel
(1909D).

A much more comprehensive scheme of classification was given by Theissen
and Sydow (1917). Here again the family Capnodiaceae is retained. The primary
divisions of the family were based on the character of the perithecium, whether
stalked or sessile, and on the colour and septation of the ascospores. In this
scheme the genera in the Naetrocymbeii (now known to be identical with the
Coccodineii) were placed in the Capnodiaceae. Here also were placed the genera
previously classified as Chaetothyriaceae, a subsection of the Microthyriaceae, by
Theissen (1913), and a fungus named by von Hoehnel Treubiomyces, which had
originally been placed in the Nectreii. According to Theissen and Sydow the
Capnodiaceae included 23 genera.

The classification of Theissen and Sydow was followed by Mendoza (Stevens,
1925) in his treatment of the sooty moulds from Hawaii.

A rather different scheme was outlined by Spegazzini (1918) in his review
of the family. The synonyms accepted by him are not always in agreement
with those advocated by Theissen and Sydow. He made the primary divisions
of the family on the character of the perithecium as did Theissen and Sydow.
Twenty genera were included in this scheme. Spegazzini considered the imperfect
fruits of the Capnodiaceae separately under different generic and specific names
as the Capnodeii Imperfecti.

Von Hoehnel (1918) again maintained that the division of the Capnodiaceae
into two families, the Capnodiaceae proper and the Coccodineae, was justified.

Von Hoehnel’s opinion was accepted by Woronichin (1925), who proposed
that the sooty mould family should be elevated to the rank of a class equal to
the Sphaeriales and the Perisporiales. For this he proposed the name Capnodiales,
and attributed to it three families: i. The Antennulariellaceae, Woron.; ii. The
Coccodineae, v. H.; iii. The Capnodiaceae, v. H. The family Antennulariellaceae
included one genus Antennulariella Woronichin, which was described as having
perithecia of the typical perisporiaceous type, in the sense of Theissen and Sydow
(1917), and the imperfect fructifications and mycelium of the Capnodiaceae.
Woronichin considered that the Chaetothyriaceae of Theissen (1913) should be
classed as an offshoot of the Capnodiales, which approach the Microthyriaceae in
the method of formation of the perithecia.

The new class Capnodiales was not accepted in the classification of Clements
and Shear (1931), and the type genus Antennulariella of the family
Antennulariellaceae was placed by them amongst the doubtful fungi.

380 THE SOOTY MOULDS OF NEW SOUTH WALES. 1,

The raising of the Capnodiaceae to the position of a class seems unjustified
at present, and investigations of the life histories of the sooty mould fungi are
necessary before even the relationships of the fungi now classed together in the
Capnodiaceae can be adequately assessed.

Woronichin (1926) proposed a_ different classification for the fungus
Pleosphaeria citri Arn. He did not agree with Arnaud that this fungus belongs
to the genus Pleosphaeria. He examined material from the herbarium of Jaczewski,
which had been identified by Briosi and Cavara, and which he maintained was
identical with that described by Arnaud. He described the perithecium as being
like a truncated cone in side view and spherical when viewed from above. The
wall is of dark brown cells and not truly parenchymatous, the ostiole is formed
by the gelatinization of the cells at the apex of the perithecium, and it is fringed
by a number of pointed bristles. Woronichin stressed the fact that hyphal threads
can be traced from the perithecium wall into the surrounding mycelium. The
spores are pale yellowish-green and 3-4 septate. His opinion was that the fungus
undoubtedly belongs to the genus Aithaloderma which is placed by him in the
Chaetothyriaceae, but which is placed in the Capnodiaceae by Theissen and Sydow.
Woronichin proposed the new combination Aithaloderma citri Woron. for the
fungus, and considered that it is closely related to A. colchicum Woron.

Gaumann (1928) accepted the classification of the sooty moulds proposed by
Arnaud. He placed them in the new family Amphisphaeriaceae of the Sphaeriales,
because of the presence of an ostiole in the perithecium of most of the members.

The classification of the sooty moulds adopted by Clements and Shear (1931)
was a modification of that of Saccardo. It was based primarily on the colour and
segmentation of the ascospores. The number of genera recognized was 34. Most
of these are superficial in habit, but several have a subcuticular mycelium.

As has been pointed out by many writers on the subject (see Clements and
Shear, 1931), the fungi classed in the Capnodiaceae show relationships both with
the Sphaeriales and with the Perisporiaceae, and in many respects appear to be
intermediate between them. ;

Caldariomyces Fumago Woron.—The fungus described by Zopf (1878) as
Fumago has been the cause of some differences of opinion. Zopf himself gave
it no specific name. Zopf grew this fungus in pure culture in solutions of various
concentrations of sugar and followed the development of a great variety of
fructifications. The simplest of these are the “Konidienbuscheln”, bundles of
conidiophores which cut off conidia at their apices. By overgrowth of the marginal
hyphae of such fructifications as these, all gradations are produced between this
type and stalked pycnidia with long or short necks according to the degree of
overgrowth. Gradations between these and short-stalked pycnidia with long or
short necks are also found. ‘Gewebefruchte”, which Zopf considered to be truly
parenchymatous in origin as compared with the previously described pseudo-
parenchymatous fruits, were also described. These are small pear-shaped pycnidia.
Zopf also found that, after some months of cultivation, yeast-like conidia were
produced by his fungus, and these under certain conditions reproduced
themselves by budding after the manner of yeasts. Other conidia were also
described.

Lindau, Arnaud, Gaumann and others have attributed the various fructi-
fications described by Zopf to the fungus Capnodium salicinum Mont. Neger
(1918) has advanced evidence to show that this is not the case. In Germany,

BY LILIAN FRASER. 381

according to Neger, the fungus described by Zopf is found only in glass houses,
whereas Capnodium salicinum is very widespread. Zopt’s fungus and Capnodium
salicinum were shown by Neger to be quite distinct. Since, therefore, the perfect
stage of the glasshouse fungus is unknown, Neger referred to it as Mumago vagans
Pers. Woronichin (1926), however, maintained that it could not belong to the
genus Fumago as described by Persoon, and proposed for it a new name,
Caldariomyces Fumago Woron. He considered that the fungus should be placed in
the Fungi Imperfecti, near Graphium. According to Woronichin Fumago vagans,
as generally known, is a mixture of Dematium pullulans and Cladosporium
herbarum. Clements and Shear (1931) did not accept Woronichin’s classification
and referred to the genus Caldariomyces as a synonym of Fumago. According
to their key, however, the fungus described by Zopf cannot be placed in the genus
Fumago which is near Alternaria.

(ii) Atichiaceae.

A fungus, Atichia glomerulosa, which is very frequently met with in European
sooty moulds (see Neger, 1918), has been the subject of much debate. Brief
reviews of the genus have been given dy von Hoehnel (1910), Cotton (1914),
and Neger (1918).

The fungus forms small round or lobed cushion-like colonies 0-5 to several
millimetres in diameter on the leaves of trees. These appear black macroscopically
and consist of radiating threads of yeast-like cells embedded in mucilage. The
inner cells are hyaline, the outer ones are darker and have thicker walls. The
cells contain numerous droplets of oil. The colonies are small and dark when
dry, but because of their mucilaginous nature they absorb water readily, and
swell at once to several times their original size.

Reproduction takes place by the formation of ascospores in asci which arise
near the surface of the colony. They may be grouped in special cushion-like
areas, or scattered throughout the thallus. Asexual reproduction takes place by
the production of many-celled conidia called “propagulae’ which are produced
in great numbers in concavities in the thallus. They consist of small branch
systems of cells similar in appearance to those composing the remainder of the
thallus, the shape varying with the species. Pycnidia of typical form have also
been described.

The genus Atichia was described. in 1850 by Flotow for the fungus known
as Collema glomerulosa Ach., which was previously placed in the Collemaceae
(Lichens). Lindau (1897) placed the fungus in the Bulgariaceae as a doubtful
genus.

A fungus of the same genus was described by Patouillard in 1904 under
the name Seuratia. Patouillard at first considered it to be a new genus of the
Capnodiaceae. Later Vuillemin (1905) in describing S. pinicola (which has
been shown by von Hoehnel to be identical with Atichia glomerulosa) made a
new family, the Seuratiaceae, for these fungi, and pointed out that they showed
affinities with the Capnodiaceae and the Celidiae. This classification was accepted
by Patouillard (1906).

Von Hoehnel (1910) pointed out that the genus Sewratia was identical with
the previously described Atichia, and placed it with the Saccharomycetes. He
placed in the genus Atichia the fungus known as Heterobotrys Sacc., frequently
found associated with sooty moulds, and described by many writers as a spore
form of members of the Capnodiaceae. The type species Heterobotrys paradozra
is now regarded as being identical with Atichia glomerulosa (Cotton, 1914).

E

382 THE SOOTY MOULDS OF NEW SOUTH WALES. i,

Mangin and Patouillard (1912), in describing a fungus belonging to the genus
Phycopsis which is closely related to Atichia, maintained that Phycopsis and
Atichia cannot properly be classed in either the Capnodiaceae or the Saccharo-
mycetes, and suggested a new group, the Atichiales, with one family, the
Atichiaceae, and two genera, Atichia Flot. and Phycopsis Man. and Pat. They
placed this group at the base of the Ascomycetes, parallel to the filamentous
Ascomycetes. They pointed out that the fungi placed in the Atichiales resemble
the Saccharomycetes in the habit of budding, and the Myrangiales in the method
of reproduction. They considered that the group was derived from the Florideae,
and compared the propagulae with the soredia of the lichens.

Gaumann (1928) retained the family Atichiaceae, describing the fungi
belonging thereto as of doubtful affinities. He considered that they may be
related to the yeasts or may be reduced Discomycetes.

Clements and Shear (1931) placed the genera of the Atichiaceae in a new
class, the Argyriales, in the family Argyriaceae. The members of this new class
are described as being characterized by extreme reduction of the apothecium.
They are said to have affinities with the Bulgariaceae, Ascobolaceae, Pezizaceae
and Myrangiaceae and are probably derived from these families by reduction.
The Argyriales is, therefore, not a natural class and contains a very heterogeneous
collection of fungi.

(iii) Fungi Imperfecti.

In 1888, Laurent carried out a number of cultural experiments on sooty moulds.
He made a great number of isolations on nutrient gelatine plates of fungi from
sooty moulds from various parts of Europe and tropical countries. The fungi
he recorded as occurring most commonly were, in order of abundance, Clado-
sporium herbarum, Penicillium cladosporioides (now considered to be the same as
Cladosporium herbarum), and Dematium pullulans. Yeasts were also obtained.
He grew these fungi in pure culture, and studied their reactions to heat and
cold. He found that in all cases a dark mycelium was ultimately formed by
each fungus (dark coloured spores in the case of the yeasts). This dark mycelium
was very similar to that forming the sooty moulds on leaves. As a result of
this and of other experiments in which he found that conidial fructifications
resembling each other were occasionally developed by the three fungi, he came
to the conclusion that these three species of fungi were forms of the one fungus,
Cladosporium herbarum. He explained this variability as being induced by the
action of the sun. The fungus Capnodium salicinum was regarded as being the
perfect stage of this supposedly very polymorphic fungus.

Dematium pullulans was first described by de Bary (1887) as an epiphyte on
the leaves of trees. In culture it produces thin-walled conidia laterally in great
abundance; these may grow by budding after the manner of yeasts. Under
adverse conditions thick-walled chlamydospores and conidia are formed. It is a
saprophyte common in all parts of the world.

Schostakowitsch (1895) carried out extensive cultural experiments on sooty
moulds, examining them particularly with regard to the conidial fructifications.
.He found three species of the Fungi Imperfecti to be associated with the formation
of sooty moulds, Cladosporium herbarum, Dematium pullulans, and Hormodendron
sp. He found no justification for Laurent’s conclusion that Cladosporium and
Dematium are expressions of the one fungus, and considered that these species

BY LILIAN FRASER.

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and Hormodendron are culturally quite distinct. Zopt’s results showing the
extreme polymorphism of Fumago were confirmed, but Schostakowitsch believed
that the yeast-like conidia ascribed by Zopf to Fumago were in reality a yeast
contaminant and not part of the life cycle.

Berlese (1895) and Planchon (1902) supported Schostakowitsch’s criticism
of the results obtained by Laurent. As the result of extensive experimental work
they found no cultural evidence of relationship between Cladosporium and
Dematium.

Arnaud (1910@) mentioned that Cladosporium and Dematium are the most
important members of the Fungi Imperfecti which go to the formation of sooty
moulds in France. He also sometimes found spores of other common saprophytic
members of the Fungi Imperfecti in sooty moulds.

In 1917 Neger published the results of his important cultural work on the
sooty moulds. He found that there are a great many more fungi composing
these moulds than was generally accepted. Dematium pullulans, Cladosporium
herbarum, Coniothecium sp., Triposporium sp., Atichia glomerulosa, Hormiscium
pinophilum were frequently found, the first two being the most important.
Helminthosporium sp., Penicillium spp., Botrytis cinerea, yeast and bacteria were
also occasionally obtained, together with the sterile mycelium of Bulgaria
polymorpha, Herpotricha nigra, Xylaria hypoxylon and others. He insisted that
it is necessary to make cultural as well as microscopical examinations of sooty
moulds in order to ascertain their composition.

Dematium pullulans has been regarded by some mycologists, notably Brefeld
(see Hoggan, 1923), as being a collective name for the imperfect form of a
number of the Sphaeriales. Amongst the species to which it has been referred
are Sphaerulina intermizta and Plowrightia ribesia. Hoggan (1923) gave a
detailed review of literature concerning Dematium, and showed that it has no
connection with the fungus Plowrightia. Bennett (19280) later obtained the
periect stage of Dematium and showed that it is unlike any of the Capnodiaceae
or Sphaeriales.

Bennett (1928a) gave a summary of evidence to support the contention
that Cladosporium herbarum and Hormodendron are not distinct species.
Schostakowitsch distinguished them by their heliotropic reactions, the nature of
the conidial wall and the type of growth in certain concentrations of potassium
nitrate, but recent workers have found no evidence for their separation.

B. Physiological.

In the earliest studies of sooty moulds there appears to have been some
doubt as to whether the fungi were saprophytes or parasites. Tulasne, for
example, did not believe that these fungi were associated with scale insects, and
pointed out that they may occur on trees which are unaffected by scale. As
later writers have shown, however, if this is the case, it is always found that
neighbouring trees are affected and that the honey-dew has dropped from them.

Although Zopf (1878) grew Fumago in culture media of various sugar
concentrations, his chief aim was to obtain stages in the life history and not
to examine the physiological requirements of the fungus. More definite is the
work of Laurent (1888) on Cladosporium and Dematium. He studied the effect
of light and temperature on these fungi and confirmed de Bary’s record that
Dematium can remain alive for long periods of drought in the form of
chlamydospores. Schostakowitsch (1895) greatly extended the work of both

384 THE SOOTY MOULDS OF NEW SOUTH WALES. i,

Zopf and Laurent. He studied conidium formation by Fumago and Dematium
at various temperatures and on various media. He found that Dematium showed
great variability. At laboratory temperatures and in media of fairly low sugar
concentration, thin-walled conidia, which grow by budding after the manner of
yeasts, are cut off. But the production of these conidia decreases with increase
in concentration of sugar in the medium, until it ceases at about 50% sugar.
After prolonged culturing at 30° C., Dematiuwm degenerates into a yeast and
reproduces entirely as such until conditions of lower temperature are restored.

The work of Hoggan (1923) showed that Dematium pullulans is very variable
in culture, and that the thick-walled chlamydospores sometimes formed by it have
remarkable powers of resistance to desiccation over long periods.

THE Soory Moutps or New SoutH WALES.

In the Sydney district the late spring and summer months are usually fairly
dry, so that during the hottest part of the year conditions do not appear to be
favourable for the growth of sooty moulds. Sooty moulds are common, however,
at all times of the year, and show a definite increase in quantity during the
winter. Sooty moulds are found on a great variety of plants affected by scale
insects, aphis, etc. In spite of the frequency of the mould, the total number of
members of the family Capnodiaceae so far known from New South Wales is not
large. It therefore seemed that it might prove of interest to investigate some
of the moulds culturally in order to ascertain whether members of the Fungi
Imperfecti and the Atichiaceae are present to the same extent as described by
Neger in the sooty moulds of Germany.

A. Fungi Present in the Sooty Moulds of New South Wales.
(i) The Capnodiaceae.

Fifteen species of the Capnodiaceae are recorded from Hastern Australia by
Cooke (1892), McAlpine (1895) and in the index of Australian fungi compiled by
Brittlebank, which is in the possession of the Council for Scientific and Industrial
Research at Canberra. These are listed below, together with synonyms and
notes on the present systematic position of the fungus.

Fungi recorded by Cooke are indicated by (C), additional species recorded
by McAlpine by (M), and further additions recorded in Brittlebank’s host index
by (B).

1. Capnodium citri B. & Desm. Victoria and Queensland (C).—Synonyms:
Capnodium citri Mont., Apiosporium citri Br. & Pass. (see Lindau, etc.), Meliola
citri Sace., Limacinia citri Sacc., Pleosphaeria citri Arnaud, Aithaloderma citri
Woronichin. This fungus does not belong to the genus Capnodium as now
accepted, and it is probably best placed in the genus Aithaloderma.

2. Capnodium elongatum B. & Desm. Queensland (C)—Synonyms: Polychae-
tella elongata Spegazzini, Capnodium Persoonii B. & Desm. (Woronichin, 1926).
Arnaud considered that this species is a doubtful one.

3. Capnodium australe Mont. Queensland (C).

4. Capnodium salicinum Mont. Eastern Australia (C).—Synonyms: Fumago
Ssalicina Pers., Apiosporium salicinum (see Lindau, etc.), Teichospora salicina
Gaumann (see Arnaud, 1910a, 1911). This is a very widely distributed fungus,
being common in the sooty moulds of all parts of the world.

5. Capnodium Walteri Sacc. Victoria (M).

BY LILIAN FRASER. 385

6. Capnodium araucariae Thuem. Victoria (B).—Synonym: Polychaetella
araucariae Spegazzini. The type species was described as an imperfect fruit,
probably of Capnodium australe.

7. Capnodium armeniaceae Thuem. Victoria (B).—This is considered to be
a doubtful species by Arnaud.

8. Capnodium callitris McAlp. New South Wales (B).—Synonyms: Limacinia
callitris Sace., Limacinula callitris Spegazzini, Phragmocapnias callitris Theissen
and Sydow. The nomenclature of Theissen and Sydow is accepted.

9. Capnodium casuarinae McAIp. New South Wales (B).—Synonym:
Microxyphiun casuarinae Spegazzini. The description of the type was based on
an imperfect fruit form.

10. Capnodium citricolum McAlp. Eastern Australia (B).—Synonyms:
Teichospora citricola Arnaud, Limacinula citricola Spegazzini.

11. Capnodium nerii Rab. Victoria (B).—Arnaud gave this species as the
pycnidial form of Capnodium (Teichospora) meridionale Arnaud.

12. Antennaria scoriadea Berk. New South Wales (C).—Synonyms: Anten-
nularia scoriadea (Berk.) Reich., Capnodium scoriadeum von Hoehnel. The fungus
found in New South Wales is best known provisionally as Antennularia scoriaded.

13. Antennaria semi-ovata B. & Br. Queensland (C).

14. Antennaria Robinsoni B. & Br. Victoria (C).—This and the preceding
species have been described in the imperfect condition oniy.

15. Zukalia loganiensis Sace. & Berl. Queensland (C).—This species is now
known as Chaetothyrium loganiense Theissen.

The most important species of the Capnodiaceae which occur in New South
Wales are Capnodium salicinum Mont. and Antennularia scoriadea (Berk.) Reich.

CAPNODIUM SALICINUM.

This fungus is common throughout the year, especially after the autumn
rains. It appears to be widespread throughout the State and is found on a great
variety of plants. The pycnidial fructifications vary considerably in size. They
are usually stout and columnar, tapering towards the mouth. The mouth is
fringed by a few stiff hyaline hair-like hyphae. Occasionally cases of branching
pycnidia are found; in some cases young pycnidia may develop from the walls
of old ones.

The pycnidiospores of Capnodium salicinum vary considerably, their appear-
ance depending on their age at the time of ejection from the pycnidium, this in
turn depending on the weather. The spores are only ejected in damp or wet
weather. The interior of the pycnidium contains mucilage, probably secreted
from the cells of the wall. In damp weather this takes up moisture and swells,
bulging out through the ostiole and taking with it any detachable spores. Outside
the pycnidium more moisture is taken up, so that the spores appear suspended
in a clear liquid drop at the mouth of the pycnidium. If this ejection takes place
when the spores are first cut off, they are oval, hyaline, single-celled, and measure
6-Su x 4-5u. But if ejection is delayed the spores may have developed further
and have become dark brown. In this condition they may be two-celled and
10-12u x 6-8u, or four-celled and 15-18u x 6-Su. Most of the spores are at the
same stage of development in the pycnidia of any one colony.

The pycnidia are produced first by the mycelium and are best developed at
about the time of the first appearance of the young perithecia. After this their
activity declines and they ultimately cease spore production, and vegetative

386 THE SOOTY MOULDS OF NEW SOUTH WALES. i,

hyphae grow out from their sides, giving them a hairy appearance. They probably
fruit for several weeks continuously.

The perithecia measure 100-150u x 90-120u; the ascospores are brown,
15-194 x 7-9u, usually three-septate laterally, with additional longitudinal septa
giving them a muriform appearance. They may be slightly constricted at the
median septum.

ANTENNULARIA SCORIADEA.

Von Hoehnel (1909c) has reviewed the species Antennularia scoriadea (Berk.)
Reich. in some detail. The type specimen from New Zealand is badly preserved,
but he examined other similar specimens from New Zealand, describing the
perithecia and renaming the species Capnodium scoriadeum (Berk.) v. H. The
pycnidial form of a species very similar to this is very common around Sydney,
but, until the perithecial fruits have been found, it is advisable that the New
South Wales fungus should be known as Antennularia scoriadea.

As a rule sooty mould colonies in which this fungus is predominant are very
thick and spongy in appearance. The mycelium is dark brown and torulose.
The pycnidia are pear-shaped, 40—-50u x 60-—70u, and produce small hyaline spores
about 2u in diameter. Antennularia occurs very frequently as a minor constituent
in moulds in which other fungi are dominant. In some such cases it may not
be found in the fruiting condition, and its presence is only revealed by platings.

Besides Antennularia scoriadea and Capnodium salicinum there are other
members of the Capnodiaceae present in the sooty moulds of New South Wales,
which produce various forms of pycnidia, but have not yet been found associated
with perithecia. These will be discussed at greater length in a later communica-
tion. Several species of Capnodium may occur in the one mould.

(ii) Atichiaceae.

A common constituent of the sooty mould flora is Atichia glomerulosa. Not
only is it found growing with sooty moulds, but it is also common alone on the
leaves of rain forest trees, and the individual colonies may attain considerable
size. From his descriptions and figures, as well as the fact that he called this
type of structure Heterobotrys, it is very probable that the glomerulae described
by McAlpine as the conidial fructifications of Capnodium citricolum and C. callitris
are colonies of a species of Atichia.

(iii) Fungi Imperfecti.

A number of types of conidia belonging to genera of the Fungi Imperfecti
have been recognized in microscope examinations of New South Wales sooty
moulds. The most frequent of these are Cladosporium, Alternaria and
Triposporium.

In order to investigate fully the Fungi Imperfecti present in moulds cultural
experiments have been carried out.

Methods.—Platings were made in the following manner. Portions of a sooty
mould of about 4 sq. mm. (less in the case of a thick mould) were scraped off
the affected leaf with a sterile needle and thoroughly ground up in a drop of
sterile water on a glass slide. A drop of the suspension of broken fragments of
mycelium and spores was transferred to a sterile Petri dish. This was then
poured with a thin layer of potato dextrose agar, and the plates were incubated
for 7-10 days at 25° C. Asa general rule two plates were poured for each sample,
and six samples were taken from each sooty mould investigated in this way.

BY LILIAN FRASER. 387
Below are given the results of investigations of three typical sooty moulds
collected in the Sydney district.
1. Host, Eugenia sp., attacked by Ceroplastes rubens.—The mould was fairly
thin and newly formed. It was collected on 19th September, 1932.

Number of Plates Percentage of
Species of Fungus. in which it Number of total number
occurred. colonies. of colonies.

Dematium pullulans ...... 12 865 57-6
Cladosporium herbarum ... 10 552 36°7
PY CA'S Came yeiyore, aceon) Susys pa eueseae ore cee 9 79 5:2
EZEMACUULUUNI SD sa tal clioetel tener cl ons 5 5 0-3
FE DIGOCCULIDESDs eaiie cis sickeictenenene 1 1 0-06

2. Host, Pittosporum undulatum, attacked by Ceroplastes ceriferus.—The
mould was thick and well established. Capnodium salicinum was present, and was
fruiting prolifically. It was collected on 20th August, 1932.

Number of Plates Percentage of
Species of Fungus. in which it Number of total number
occurred. colonies. of colonies.

Dematium pullulans ........ 12 820 51-2
Cladosporium herbarum 12 394 24-6
Capnodium salicinum ...... 9 282 17-6
IPOMCOUROTE BUS Sa ododoopbos 6 36 2-2
IVSC RIS Cama etrenc ns eau isvcysueer cts susie (mie 5 25 1.06)
FAN LETIVGTEMM ISDS | ciaretalclehelel« «ere 10 24 1-5
ASWOMSGS B05 “Sauoceuotauno0e 2 2 0-12
TH WCOGCOND Gs soocouncbous 5 12 0-75
(QUAGIAS it Bs Bee Be eae cr cere 4 4 0-25

3. Host, Arbutus unedo, attacked by Ceroplastes rubens—Antennularia
scoriadea was present. The mould was collected on 1st June, 1932.

Number of Plates Percentage of
Species of Fungus. in which it Number of total number
occurred. colonies. of colonies.

Cladosporium herbarum .... 12 325 55:8
Dematium pullulans ....... 12 0s 18-5
Antennularia scoriadea : 9 48 8-2
BYICAIS Hag cwaraus/ soccer eres aictevedovatocd<iorees 10 42 7-2
ASIOMSIG Son U4 mata pecgasoc me 26 4-4
ANMGAPRUCHUR, “GOS “Ae ceo boos euc 5 14 2-2
ASDOOMSIG B13 séaceccadcoe 2 a 0-86
IPOTHCOMMH BI “cbascbnoon66 2 3 0-5
IFTKCOCCOMO GI cocococcoodc 2 3 0-5
ORES arses bos seSeie swore 6 8 1°3

About 100 such investigations were carried out. It was found that some
of the fungi, e.g., Penicillium spp., were only found occasionally, and in relatively
small numbers. They therefore could not be regarded as permanent constituents
of the sooty moulds. Others which occurred still less frequently, and were found
perhaps only once in the course of an investigation, e.g., Fusarium sp., were
considered to be purely accidental.

388 THE SOOTY MOULDS OF NEW SOUTH WALES. 1,

The percentages of the members of the Capnodiaceae shown by this method
are often far short of the percentage expected from a microscopical investigation
of the sooty mould. This is probably due to the fact that they grow very slowly
in culture, and are frequently covered by the faster growing members of the
Fungi Imperfecti.

The fungi found to be the commonest and most widespread are: Dematium
pullulans, Cladosporium herbarum, Asbolisia spp., Alternaria spp., and
Triposporium sp.

Dematium pullulans.—This fungus is present in greater or less degree in all
the moulds so far investigated, and in some cases seems to form practically pure
colonies. The mycelium is dark and more or less torulose. The conidia are
difficult to identify microscopically as they resemble the resting spores of dark
coloured yeasts. A number of different strains have been obtained, varying chiefly
in the colour of the mycelium when grown on a standard agar.

Cladosporium herbarum.—This fungus is often found in association with
Dematium, but under certain circumstances it may also form practically pure
colonies. It is present in a great percentage of the New South Wales sooty
moulds. Several strains, differing in the habit of growth on a standard agar,
have been obtained.

Asbolisia spp.—These fungi are less common than Cladosporium and Dematium.
They are characterized by having small round dark pycnidia and small hyaline
spores.

Alternaria spp.—These fungi are widespread but never very abundant in
sooty moulds. Those most frequently found are characterized by having a very
dark coloured mycelium.

Triposporium sp.—This fungus is often present in sooty moulds. It is not
so common as Alternaria but is widespread, and when present is very abundant
and fruits profusely. It is easily identified by its large star-shaped conidia.

Other species of Fungi Imperfecti which can only be detected in culture and
are much less frequent are: Brachysporium sp., Penicillium spp., Epicoccum spp.,
Fusarium spp., and yeasts, including pink, white and black coloured forms.

B. Development and Distribution.
It has been found in the course of the work that there are two types of
sooty moulds present in the Sydney district, (i) perennial, and (ii) annual.

(i) Perennial Moulds.

The perennial type is by far the most common, and represents the highest
development attained by the sooty mould community. A member of the
Capnodiaceae is always dominant. These moulds re-establish themselves very
rapidly if partly washed away by heavy rain or after flaking off the host
leaves in dry weather. They are found exclusively on perennial plants.

(ii) Annual Moulds.

The annual type of mould is found on herbs which have been attacked by
aphis. These moulds are commonest in damp weather, and consist exclusively
of members of the Fungi Imperfecti, the dominant fungus being usually Dematium
or Cladosporium.

The development of the sooty mould has been followed in a number of cases
by selecting leaves showing various stages in the appearance of the mould from
the earliest traces of mycelium to the final stage. Plate cultures have been

BY LILIAN FRASER. 389

made from these leaves. Specimens of perennial and annual moulds have been
examined in this way.

The first steps in the development are quite similar in each case. The nature
of the pioneer fungus varies, but Dematium is by far the commonest. Occasionally
the pioneer fungus may be a Capnodium, but usually this is only the case if the
mould is developing on a shrub close to others which support a sooty mould in
which a Capnodiwm is dominant. The growth of these first fungi forms a light
network over the surface of the leaf, and this forms a matrix which catches and
holds other fungal spores, and forms a ground mass in which they may germinate.
Cladosporium usually appears quite early in the development of the mould.

The subsequent development depends largely on the environmental conditions.
If the neighbouring moulds are of the annual type, or if there are no moulds
in the vicinity, the new sooty mould will be composed of Fungi Imperfecti for a
considerable period. In most cases, if the mould is on a perennial plant, develop-
ment continues, so that after a longer or shorter period the final stage is attained
This is effected by the appearance of a Capnodium which ultimately becomes the
most important fungus in the mould. Other Capnodiums may also appear, and
the mould takes on its perennial form. The time necessary for such a change
depends on the chances of infection with a Capnodium.

In a few of the cases examined this development was not completed. Here
fairly thick moulds had been formed by the growth of Dematium pullulans, with
Cladosporium and yeast also present in small amounts. The mould was perennial
and similar in outward appearance to those in which Antennularia scoriadea was
dominant, but the luxuriance of the growth of the Fungi Imperfecti made it
impossible for members of the Capnodiaceae to become established. Strong growth
of Dematium appears, therefore, to be inimical to the growth of Capnodium.

In the case of annual plants the sooty mould formed may become fairly
thick, but in all the cases examined it was composed of the annual type of fungus
association. The annual mould may be taken as a stage in the development of
a perennial mould, in which there is not sufficient time for a Capnodium to become
established.

In the case of sooty moulds developing on a perennial plant in the vicinity
of moulds in which a Capnodium is present, it has been found that development
proceeds immediately to the formation of a mould of the perennial type without
any intervening stage.

The members of the Capnodiaceae grow well together and seem to have no
mutually antagonizing effect. On the other hand, as has been shown, a growth
of Dematium, Cladosporium and yeasts forms a combination into which it is difficult
for a Capnodium to enter. It has been found also that a strong development
of Alternaria, Asbolisia and Penicillium considerably reduces the amount of
Dematium and, to a less extent, Cladosporium present.

The fungi obtained from culture from the moulds can be classified into three
groups: :

1. The Perennials.—These are the chief constituents of the perennial moulds.
They are characterized by having a dark, resistant mycelium which can withstand
a considerable amount of desiccation, and can commence growth whenever con-
ditions are favourable. The fungi included in this class are the members of
the Capnodiaceae, Atichia glomerulosa, Dematium pullulans, Cladosporium
herbarum and possibly Triposporiumn.

THE SOOTY MOULDS OF NEW SOUTH WALES. i,

Go
is}
(=)

2. The Ephemerals.—These are the fungi other than Dematium and
Cladosporium which make up the bulk of the annual moulds, and appear only in
favourable weather. They decrease very considerably in amount during hot and
dry weather, and reappear when climatic conditions are suitable. They evidently
cannot withstand insolation to the same extent as do the members of the first
class, and rely chiefly on their spores for tiding over adverse periods. This class
is represented by Alternaria, Brachysporium, Asbolisia, Epicoccum, yeasts and
possibly Penicillium.

3. The Accidentals.—These are fungi which develop from spores blown by
chance on to the mould, and which grow only in the most favourable weather,
dying off completely in hot dry periods. They never form an important part
of the mould. They include Fusarium spp., Mucor, Aspergillus, bacteria, etc.

THE PROBLEM OF POLYMORPHISM.
A. Historical Review.

As pointed out in a previous section, the fact that a number of fungi may
grow intermingled and fruit concurrently in a sooty mould, leads at once to the
idea that there is a single polymorphic fungus concerned in its formation.

Tulasne assigned many imperfect forms to Capnodium salicinum$ most of
which were probably members of the Fungi Imperfecti.

Thuemen (1890) also spoke of the extraordinary diversity of the spore forms
of the ‘‘Russtaupilze”’. Penzig included as an imperfect stage of Meliola camelliae
the colonies of Atichia associated with it. The work of Zopf on a rather poly-
morphic species gave such ideas much support. Laurent went to the extreme of
trying to correlate a number of distinct species of the Fungi Imperfecti culturally,
suggesting that they were identical, and the imperfect stages of Capnodium
salicinum.

Schostakowitsch (1895), Berlese (1895), and Planchon (1902) did much to
establish the fact that the more common conidial forms associated with sooty
moulds are separate fungi.

When McAlpine described Capnodium citricolum (1896a), he made a special
study of the spore forms associated with it. That he was alive to the possibility
of other fungi being involved in the production of the mould is clear, since he
said (p. 470): “In order to prove the fact of polymorphism it would be necessary
to sow pure cultures and watch the development of the different forms under
strictly test conditions, for otherwise the forms found together might be really
different, and constitute merely a case of association.” He described seven
imperfect forms for C. citricolum and five for C. callitris (1896b).

Von Hoehnel (19090) also spoke of the diversity of the conidia produced by
members of the Capnodiaceae, which, he said, include Torula, Triposporium,
Helminthosporium, etc., together with many kinds of pycnidia which are often
vertically elongate.

Vuillemin (1908) pointed out that Bernard had described and figured Sewratia
(i.e., Atichia) in the life histories of Capnodium javanicum Zimm. and C. stellatum
Bern. from Java.

Arnaud (19100) also considered that Atichia was a stage in the development
of the Capnodiaceae.

The more recent workers on the sooty moulds are inclined to limit the
number of spore forms attributable to the members of the Capnodiaceae. Both
Woronichin (1926) and Boedijn (1931) spoke of Triposporium and other spore

BY LILIAN FRASER. 391

forms as being associated with these fungi, but considered that they were probably
distinct from them. Boedijn has, however, reported that he has observed yeast-
like budding of the mycelium of Chaetothyrium and Capnodium.

Gaumann (1928) spoke of the great diversity of the conidial fructifications
of the Capnodiaceae.

Similarly, imperfect conidial and pyenidial forms have been attributed to the
species of the parasitic genus WMeliola and related fungi, but it has been shown
by Stevens (1918) that these are separate fungi associated with Meliola and are
in most cases probably parasitic upon it.

In the same way it is probable that in all cases the spores of genera of the
Fungi Imperfecti which have been found associated with the Capnodiaceae are
produced by associated fungi, and that the only imperfect fruits produced by the
Capnodiaceae are pycnidia, and that these are always specific.

B. Cultural Evidence.

The strongest evidence that a variety of pycnidial forms may be attributed
to the one sooty mould fungus is provided by the researches of Zopf. It must
be pointed out, however, that the fruits described by him, except for a few
yeast-like forms which were probably contaminants, all produced spores of the
same size and shape. Moreover, all the types of fructifications were related by
a large number of intermediate fruit bodies, and could be placed in a definite
series. The various pycnidial fruits which were described, for instance, for
Capnodium citricolum by McAlpine bear no such relation to one another, and
produce spores of greatly varying size and shape.

The fructifications described by McAlpine for Capnodium citricolum are
reviewed below:

1. Coniothecium.—A fungus which produces spores similar to this has been
isolated by Neger (1918) and is quite distinct from Capnodium. This fungus,
however, has not yet been isolated from New South Wales sooty moulds and may
not occur here. On the other hand, the Coniotheciuwm-like gemmae described by
McAlpine are quite common. In some cases these gemmae constitute practically
the whole of a light mould. To determine the nature of these fungi, cultures
have been made, and in every case Dematium pullulans or a black yeast is
obtained. These two fungi form very similar gemmae.

2. Glomerulae—As has been previously pointed out, these are probably
colonies of an associated fungus, Atichia.

3. Triposporium.—Spores similar to those described have been found many
times. Cultures of these spores have been made from a diverse number of sooty
moulds, and in all cases gave rise to pure cultures of an imperfect fungus bearing
Triposporium spores alone.

4. Spermogonia.—These are short pycnidia with small rod-like spores.

5. Antennaria.—These are pear-shaped or oval pycnidia with small round or
oval spores.

6. Ceratiopycnidia.—These are elongated or cylindrical pycnidia producing
oval, single-celled, hyaline spores.

7. Pycnidia.—These are also elongated and cylindrical, and produce spores
which are one to three septate, and hyaline or brown.

Pycnidia of all these types have been found and cultures made from the
spores produced by them. In each case the fungus obtained from one type of
pycnidium differed essentially from the fungi obtained from the pyenidia of other

392 THE SOOTY MOULDS OF NEW SOUTH WALES. i,

kinds, and it is certain that a number of different fungi are involved in the
production of the diverse types described as belonging to the one fungus by
McAlpine.

Spore cultures from the “pycnidia” yield cultures which are identical with
ascospore cultures of Capnodium salicinum. It is, therefore, certain that these
fructifications do not form part of the life cycle of C. citricolum.

The small “Antennaria” pycnidia are referable to Antennularia scoriadea.

Two types of pycnidium, therefore, remain unaccounted for, the Spermagonia,
and the Ceratiopyenidia. Of these, it is probable that one is really the imperfect
form of C. citricolum, but until ascospore cultures can be made, it is impossible
to decide which.

To test the hypothesis of polymorphism further, spores of Alternaria and
Cladosporium were cultured directly from a diverse number of moulds. In all
cases pure cultures of Alternaria and Cladosporium were obtained. No other
fruit bodies were produced by these fungi in culture.

To date no evidence has been obtained culturally that any of the Capnodiaceae
produce conidia of any kind, or pycnidia of more than one kind.

CONCLUSIONS.

The results given above indicate that the New South Wales sooty moulds
show resemblances, on the one hand, to sooty moulds of tropical regions, in that
several species are commonly found contributing to the formation of a single
mould. On the other hand, there are resemblances to the sooty moulds of the
cool north temperate regions. Such saprophytes as Cladosporium herbarum,
Dematium pullulans and Atichia glomerulosa are common and widespread in
New South Wales, as well as in the north temperate region. The absence from
the New South Wales sooty moulds of Botrytis cinerea, bacteria, and many fungi
recorded by Neger is noteworthy. In this respect the local moulds seem to be
intermediate in type between those of the tropical and those of the temperate
regions.

The habitat of the mould is a peculiar one, and the fungi which thrive in
it must possess modifications or properties which enable them to exist under
conditions which show a great range of temperature, light intensity and humidity.
They must be able to utilize casual moisture such as rain and dew, as well as
honey-dew, for growth, and remain unhurt through periods of great insolation.

In the temperate parts of the earth sooty moulds are best developed in moist
cool weather, when the light intensity is less than maximum. These conditions
are favourable for the growth of fungi generally, and it is therefore not remarkable
that the sooty mould should include the common saprophytes of decay.

On the other hand, tropical sooty moulds live under atmospheric conditions
differing from those obtaining in temperate countries. Temperature, light intensity
and humidity all have a high value, and the result is that there is a great variety
of specialized fungi present, chiefly members of the Capnodiaceae, which are
restricted entirely to the honey-dew covered parts of plants for their development.

ScaLE INSECTS CONCERNED.
The following important scale insects associated with the formation of sooty
moulds in New South Wales have been kindly determined by Mr. Froggatt.
Ceroplastes ceriferus on Pittosporum, Bursaria, Laurus, Citrus spp., and many
other plants.

BY LILIAN FRASER. 393

Ceroplastes rubens, particularly on Hugenia spp., but also very common on
Bursaria, Citrus and many other plants.

Lecanium hesperidum, on Bursaria, Citrus, Castanospermum, ete.

Tachardia melaleucae, on Leptospermum spp.

Others with which sooty moulds have been found associated are: Hriococcus
eucalypti, on Bursaria spinosa; Ctenochiton eucalypti, on Angophora spp.;
Dactylopius albizzia, on Acacia discolor; Eriococcus leptospermi, on Leptospermum
spp.; and Mytilaspis crassi, on Jacksonia sp.

The identity of the scale insects associated with the mould appears to exert
little influence on the species of fungi which occur. Similar moulds may develop
on the secretion of different species of scale insects as far as conditions of
environment permit.

SUMMARY.

1. A brief historical account is given of work done on the composition,
systematics and physiology of sooty mould fungi. Sooty moulds usually consist
of a number of different fungi growing together to form a community. These
fungi belong to the following groups: (a) Capnodiaceae, which includes the most
conspicuous and important members of the flora; (0) Atichiaceae, the commonest
species of which, Atichia glomerulosa, is a widespread constituent of sooty moulds;
(c) Fungi Imperfecti, which may in some cases be present to the exclusion of all
other kinds of fungi. Of these, Dematium pullulans and Cladosporium herbarum
are the most important.

2. A review of the species of sooty moulds reported from New South Wales
is given.

3. It has been found that in New South Wales the sooty moulds Capnodium
Salicinum and Antennularia scoriadea are the commonest members of the
Capnodiaceae present. Atichia glomerulosa and the Fungi Imperfecti, Dematium
and Cladosporium, are widespread.

4. There are two types of sooty moulds in New South Wales: (a) Perennial
moulds which develop on perennial shrubs and trees; (0b) annual moulds which
develop on annual herbs attacked by aphis, and which often precede the perennial
moulds on trees and shrubs. The perennial mould consists of members of the
Capnodiaceae, together with Atichia and a variety of the Fungi Imperfecti. The
annual mould consists largely of Dematium, Cladosporium, Alternaria, Asbolisia,
Triposporium, yeasts, etc.

5. Dematium is the pioneer fungus of the sooty mould flora in most cases.

6. It is shown that all the spore forms attributed by McAlpine to C. citricolum
do not belong to the one species.

The writer wishes to express her thanks to Professor T. G. B. Osborn, of the
Department of Botany, Sydney University, for suggestions and helpful criticism
throughout the course of the work, and to Mr. W. W. Froggatt, late Government
Entomologist of New South Wales, who kindly identified the scale insects found
associated with sooty moulds.

While this paper was in the course of preparation for the press the writer
was fortunately able to see, through the kindness of Miss HE. E. Fisher, M.Sc., of
Melbourne, an advance proof of a paper by her: ‘On the ‘Sooty Moulds’ of some
-Australian Plants’, which is to appear in Proc. Roy. Soc. Victoria, 45 (N.S.),
Pt. 2, 1933. Miss Fisher considers that the fungi Capnodium citricolum McAIp.,
and C. Walteri Sacc., are identical with C. salicinum Mont. She follows the nomen-

394 THE SOOTY MOULDS OF NEW SOUTH WALES. i,

clature of Arnaud and Gaumann, referring the fungus to the genus TJeichospora.
Miss Fisher’s conclusions will be discussed at greater length in a later
communication.

Literature Cited.

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—_———,, 1910a.—-Atichia Treubii v. H. (Saccharomycetes). Ann. Jardin Botan.
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LINDAU, G., 1897.—Perisporiales. Die Naturlichen Pflanzenfamilien, Engler und Prantl,
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BY LILIAN FRASER. 395

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Abstracts only of papers marked thus were available to the writer.

NOTES ON THE AUSTRALIAN SPECIES OF THE FAMILY PAUSSIDAE
[COLEOPTERA].

By the late T. G. SLOANE.
(Prepared for publication by H. J. Carrer, B.A., F.E.S.)

[Read 25th October, 1933.

The late T. G. Sloane was greatly interested in the Paussidae and, inter-
mittently, gave much close study to it, at least from 1920 to 1927. Among papers
left by him were four note-books (together with a photo of Westwood’s plate)
containing detailed results of this study, though the notes had not been collected
into a coherent whole. Up to the year 1924 two authors, Westwood and Macleay,
had described the great majority of recorded Australian species. Unfortunately,
two publications by these authors on the subject clashed: and no entomologist has,
so far, tried to disentangle their probable overlapping. Sloane had clearly arrived
at determinations of the greater number of these species, had classified them into
groups, and, further, had tabulated the species in these groups, when a new author,
Professor Hermann Kolbe, in 1924, published four papers on Australian Paussidae?’.

In these papers he described some 29 species as new and arranged the genus
Arthropterus in two main groups which he again subdivided into eight subgenera.
The greater part of the subsequent notes by Sloane was devoted to an attempted
elucidation of Kolbe’s work, and a determination of the species described by him.
A complete set of these four papers by Kolbe formed one of the four note-books
mentioned above. Sloane’s note-books are now lodged in the library of the
Linnean Society of New South Wales. Certain of his conclusions seem worth
publishing, so that future students of the family may benefit by the work of one
of the most careful and accurate of Australian Coleopterists.

The family Paussidae is strikingly different from other Coleoptera. Sluggish
in movement, in general with greatly widened appendages, they are, in other
parts of the world, usually associated with ants.or termites. In a few instances
they have been thus observed in Australia. Thus Sloane quotes a letter from
C. Oke: “One from Eltham taken in nest of Iridomyrmex rufoniger. The pair
from Bendigo were obtained under a fairly large stone, well imbedded, in a nest
of Campanotus clavipes, being the only ones I have found in association with
ants.” Lea reports A. brevis in nest of Hctatomma metallicum, and A. angulatus
Macl. taken at Bowen by A. Simson in ant’s nest. Mjéberg found A. picews West.
on trunks of trees at night. Westwood records A. hopei as found at Port Phillip
under bark and dried cowdung. Sloane says: “I have only found Arthropterus

1 Westwood in Thesaurus Ent. Oxon., 1874, and Macleay in Trans. Ent. Soc. N.S.W.,

2 (a) Die Australischen Paussiden der Wassmann’schen Sammlung (Tijd. v. Ent.,
1924); (b) Uber einige Paussiden Arten Australiens (l.c.); (c) Australische Paussiden-
Arten in deutschen Museen (Ent. Mitteil., 1924): (d) Zur Kenntnis der Paussiden
Australiens: untergattungen von Arthropterus (Deutsch. Ent. Zeit., 1924).

BY THE LATE T. G. SLOANE. 397

under logs or at light in the evening.’ In my own (H.J.C.) experience all the
species captured have been found under bark, logs or stones, without apparent
association with ants or termites. They are often concealed by their colour
likeness to the background or by the inequalities of the surface of logs. I have
found the commoner Sydney species, A. brevis, in small colonies under bark of
Eucalyptus. The rarity of these insects constitutes a difficulty. Sloane notes a
number of species in which only one sex was known to him, although he had
probably more material for study than any former Australian student of the family,
since many collectors, including myself, had either given or lent their specimens,
and he had studied closely the collections in the chief museums of Australia—
especially the Macleay types. He also, through the courtesy of Mr. H. E. Andrewes,
who compared specimens sent with Westwood’s types, obtained much essential
information and copious notes on these.

Structure.—Sloane redescribed the genus Arthropterus, also several of the
older species, including the genotype A. Macleayi Don. In Arthropterus the
antennae, apparently—and hitherto considered as—10-segmented, are shown to be
11l-segmented in all recorded species, the 2nd segment being a nodule hidden in
the swollen scape. The presence of this very small 2nd segment was noted by
Dalman for the genus Hylotorus (as described by Westwood). Sloane says: “I
have clearly seen it in Platyrhopalus and Ceratoderus.”

The structure of the underwings is of interest in view of the accepted idea
that throughout the family the underwings are always well developed (cf. Desneux
in Wytsman’s Gen. Ins., 1905).

The species are found to be (a) fully winged in both sexes, (b) winged in dg,
flightless in 2, or (c) flightless in both sexes, as follows:

(a) g winged, 9 wings large, though reduced: A. wilsoni, A. piceus.

(b) ¢ winged, 2 wings rudimentary: A. denudatus, A. waterhousei, A. ovicollis,
A. mastersi. A. rockhamptonensis.

(c) g and 9 wings rudimentary: A. hopei, A. brevis, A. westwoodi.

Many other structures are noted in great detail, no single anatomical feature
being omitted from a meticulous study that includes many measurements,

Classification.—Except for the single species of Paussus (P. australis Blkb.)
from North Queensland, which Sloane had not seen, the Australian Paussidae are
all contained in the two genera Megalopaussus and Arthropterus, differentiated
thus:

Antennaesevidently, Ti-sezmenteds ayasiccce ici eicieheielsicie oi ciel eielers oe oie) veiw ete «ic ene Megalopaussus
Amtennacwapparcntlysnlli0-Sesmenteduace yj cinaniereicie cisicee cs ceicicichel</cicliesie oa careles Arthropterus

Megadlopaussus amplipennis Lea.—Sloane writes: “I examined the type and
could fix on no point of difference between it and Arthropterus except the visibility
of the small 2nd segment of the antennae. I do not think it has as much relation-
ship to Protopaussus as to Arthropterus.”

The following is Sloane’s list of Australian Paussidae with synonyms
(excluding Kolbe’s species) and with known localities.

List of Australian Paussidae (apud Sloane 22/12/23).
ARTHROPTERUS
angulatus Macl. Rockhampton, Q.
angulicornis Macl. Ipswich, Q.
articularis Elst. S. Aust.
brevicollis Macl. N.S.W.
brevis West. N.S.W.

398 AUSTRALIAN SPECIES OF THE FAMILY PAUSSIDAE,

cylindricus Mast. (= subcylindricus West.; attenuatus Gestro). Aust.

darlingensis Macl. N.S.W.

denudatus West. (= angusticornis Macl.; kingi Macl.; politus Macl.; latipennis
Macl. 2). Q., N.S.W., S. Aust.

depressus Macl. Tweed River, N.S.W.

elongatulus Macl. Gayndah, Q.

foveicollis Macl. Sydney.

foveipennis Blkb. N. Terr.

hirtus Macl. N.S.W.

hopei West. (= picipes Macl.; subampliatus Macl.; montanus Macl.). S. Aust., N.S.W.

howitti Macl. Vict.

howittensis Mast. (= howitti West.; dohrni Kolbe). Vict.

macleayi Don. (= bisinuatus Macl.). N.S.W.

mastersi Macl. (= cylindricollis Macl.). Gayndah, Q.

nigricornis Macl. (= humeralis Macl.). Q., N.S.W.

occidentalis Blkb. W. Aust.

ovicollis Macl. S. Aust.

parallelocerus West. (= turneri Macl.). Lane Cove, N.S.W

piceus West. (Phymatopterus). (= latus Lea; macleayi West.; distinctus Thoms. ;
ceraptoides Mjob.). N.S.W., S.Q.

punctatissimus West. S. Aust.

riverinae Macl. N.S.W.

rockhamptonensis Macl. Q.

subsulcatus West. Aust.

waterhousei Macl. S. Aust.

westwoodi Macl. (= adelaidae Macl.; odewahni Macl.; puncticollis Macl.; scutellaris
Macl.; subcylindricus Macl.; wyanamattae Macl.; melbournei West.). Q.,
N.S.W., S. Aust., Vict.

wilsonit West. (= neglectus Lea 9; quwadricollis West.). S. Aust., Vict., N.S.W.

MEGALOPAUSSUS Lea.
amplipennis Lea. Kuranda, Q.
Paussus L.
australis Blkb. N.Q.

Australian Paussidae described (or noted) by Kolbe.

Arthropterus: ambitiosus Kolbe (Aust.), brunni Kolbe (Aust.), daemelianus Kolbe
(Peak Downs, Q.), discrepans Kolbe (Moreton Bay), dissidens Kolbe (Aust.),
donovani Kolbe (West. MS.) (Port Denison, N. Q’land), eruditulus Kolbe (Gayndah,
Q.), fraternus ? Kolbe (Aust.), geminus Kolbe (Q’land), horni Kolbe (Port Darwin,
N. Aust.), insidiosus Kolbe (Aust.), limitans Kolbe (Aust.), moretoni Kolbe
(Moreton Bay, Q.), negligens Kolbe (Peak Downs, Q.), novellus Kolbe (Aust.),
ominosus Kolbe (Aust.), pellaw Kolbe (Aust.), pervicax Kolbe (Q.), petax Kolbe
(Gayndah, Q.), refectus Dohrn (Aust.), schismaticus Kolbe (N. Aust.), schroederi
Kolbe (Moreton Bay, Q.), simiolus Kolbe (Aust.), socius Kolbe (Aust.), spadiceus
Kolbe (Aust.), sphinw Kolbe (Dohrn MS.) (Peak Downs, Q.), subangulatus Kolbe
(Aust.), suspectus Kolbe (Rockhampton, Q.), wasmanni Kolbe (Peak Downs, Q.).

From the above it will be apparent, as noted by Sloane, that some of the
species have a very wide distribution.

The species of Arthropterus were arranged into seven groups, denoted by
type species (1) articularis Hlst., (2) denudatus, (3) parallelocerus, (4) hopei,
(5) wilsoni, (6) macleayi (to include westwoodi), (7) piceus (to include
brevis).

“In arranging the species of the genus Arthropterus sensu lato’, Sloane
writes, “I have preferred to use species groups rather than the subgeneric names
suggested by Westwood and Kolbe.” These were tabulated as follows:

Group tabulation.

1 (4) Antennae narrow, 4-9 less than 2% times broader than long; segment 8 equal,
legs narrow.

2 (8) Antennae, 3-10 moniliform, turbinate, each segment not widely applied to
preceding; apical sepment oval ..................02e000- articularis group

BY THE LATE T. G. SLOANE. 399

3 (2) Antennae with flagellum flat, strap-like; 38-10 applied to preceding by full
Wakolio k Gyoptoenl, Cefehoakepous Cholop=! Goa nnonoonndoandoenanododdoc denudatus group

4 (1) Antennae wide, 3-10 not less than 3 times broader than long, segment 3
subequal or unequal.

5 (6) Antennal basal segment of flagellum subequal (scape not transverse, rarely
with post-angles prominent, post-femora evenly rounded on anterior margin
EME MORN ROR OT NRC OM ICAL srousiey orcionecbepeleveueds (sushetets, tousie levis ioiecevetels parallelocerus group

6.(5) Antennal basal segment of flagellum unequal, angles of posterior side triangu-
larly prominent.

7 (10) Post-femora evenly rounded on anterior margin.

Sat CD)) Mer: OUMOLAXamt el aG OMe LuMUL UTM LD CELLS trre,ta)re) csclloifel/elcyollsi cllniielieliclie, ejiailel eel silaiietetstiele)icie) <i (e hopei group

9 (8) Prothorax wide (subquadrate, strongly, roundly narrowed to apex), basal
angles distant from pedicel and strongly reflexed, tarsi elongate, segment 2
about twice as long as broad, scape of antennae columnar (longer than
DLEACth era tet DOR ew ye uater sie cme reyes, ote ai cleies aleve era) alaneatenate meters wilsoni group

10 (7) Post-femora wide, anterior margin strongly bent about basal fourth, thence
oblique to base.

11 (14) Prothorax and elytra strongly punctate (also head).

12 (13) Form elongate, prothorax evidently longer than broad ........ macleayi group
13 (12) Form stout, prothorax usually broader than long, sometimes length and breadth
subequal.
14 (11) Prothorax and elytra laevigate (only microscopically punctate) ..............
Sood odDOODS SUS DODO OOO RONVHON OO UODOHOODOMUONHONODOOD OOOO GODE piceus group
Of these groups, articularis Elst. stands alone; the piceus group contains only
brevis Westw., besides the group type, from which it is readily distinguished.
Of the other groups I can only find three completely tabulated, as follows:
parallelocerus group.

1(4) Elytra more than twice longer than head and prothorax together ............
Ie TO fee erat enc ti ae ta ee Menem alos Sf, Shcubas fal atone sinenaneits planicornis, nN. sp.

2 (3) Antennae narrow and parallel, segments 5-10 three times broader than long,
flagellum seen from side thin, apical flat beneath .............. brevicollis

3 (2) Antennae wide and parallel, segments 5-10 four times broader than long,
flagellum seen from side moderately thick, apical convex beneath .. howitti

4 (1) Elytra not twice as long as head and prothorax together.

Dm (6) poe Othorax depressed. on) disk ((subquad))) i055 sss se rockhamptonensis

6 (5) Prothorax convex.

7 (8) Antennae thin, segments of flagellum four or five times broader than long, flat
ID SINCE Ce rawnct suctiorcy stoner eb tek Neuere cuetadet Meet onar ec enialan side <eGisuea le Subme che Newnes species

8 (7) Antennae stouter, segments of flagellum four or five times broader than long,
convex beneath.

9 (10) Head coarsely punctate, antennae with segments of flagellum four times broader
CATs OM Syeted pacastee ewes cre oer sense eas ateeebeks [ses ofa essex eoet en ens sustehe ar ¢,/eeiat oie parallelocerus

10 (9) Head finely punctate, antennae with segments of flagellum five times broader
Hee OT Seer eyerte se crea cue eee Nene eR eaah crahic entroniever oyeceetay le ai aivehenolts Mudgee species
The species from Newnes and Mudgee were species proposed as new, but are not
described.
hopei group.

1 (10) Hirsute species. Setosity long and erect.

2 (3) Orbital tubercle thick and prominent, post-tarsi elongate, segment 2 twice as
long as broad (2), post-trochanters pointed. Length 14 mm. .... ovicollis

3 (2) Orbital tubercle not very prominent, post-tarsi stout, segment 2 little longer
than broad (@), post-trochanter obtuse or truncate.

4 (7) FPost-tibiae narrow, not or scarcely wider than post-femora.

5 (6) Antennae elongate—segments of flagellum three times as wide as long, apical
segment large, broader than long, not narrower to base, head punctate but
neck smooth, prothorax with large separate punctures, post-trochanters
truncatewatnavpe xs Maenothe mimi —d Siti.) -apetaces ciechcies ele seis ee waterhousei

6 (5) Antennae short—segments of flagellum four times as wide as long, apical

segment roundly narrowed to base, head and neck densely punctate,
prothorax closely punctate, post-trochanters cordate, obtuse at apex.
Ib(ak=Ad oly, TUES sagbaalcrers nasa eee SO. 0:0 CCC Eee ROME TERE nO ROG eae ene Victorian species

400 AUSTRALIAN SPECIES OF THE FAMILY PAUSSIDAE,

UG) Bost-tibiaerwidermthanteremorart scarey tien ieiedetee eier-iesei ren tet-u-renatts New species*
8 (9) Antennae with segments of flagellum three and a half times as wide as long,
scapeusubtransverse. wuens thy 925s marae eee cpeicteiensieiclen sreiiicieieicieeiereuenons hopei
9 (8) Antennae with segments of flagellum four times as wide as long, scape wide,
inner apical angle prominent, obtuse. Length, 12 mm. ...... nigricornis

10 (1) Setosity short (post-trochanters cordate).

i Gl2)eProthoraxyborderedaty middle ot isidestuens cher l2) srr eerie) ones eiiallelelialet-lonodaielele
PEI CHOU ETRE CE ERCER CLS Cl REA A CHCA CRECIERE CRUE Oe CoRe ecient coe Otome ae G othe cylindricus Mast.

L2G) ProthoraxswithoutmlateralmpoLrdenurarierstaacr reine merch orcichersieneneions mastersi

macleayi group, westwoodi subgroup.
1 (2) Outer angle of scape of antennae not prominent, post-femora with anterior
side evenly arcuate, prothorax bifoveate behind apex .......... foveicollis
2 (1) Outer angle of scape prominent, post-femora with anterior side uneven, broad
and strongly oblique to base.
34) ee VMietasternumylare ene (cpmawineed)) i crsirenwemcenscherncnonn heleetneu rein ienstcnerenene occidentalis
4 (3) Metasternum small. a
5 (6) Prothorax not short, of variable width, broader than long, lateral border
narrow, not explanate at basal angle, these marked ............ westwoodi
6 (5) Prothorax transverse, lateral border explanate at basal angles, these obtuse
Silene Wibcai get auveiranss eee Pe sue Maa et ica a oe sad lve re Nese eNa PENG Lee Sksuanetede ld iemetrenateys cribrosus, Nn. sp.

In a later table Sloane included angulatus Macl. as allied to 6 above, but
distinguished by “prothorax decidedly narrowed to base’’.

Kolbe divided Arthropterus into two main divisions on the form of the
prothorax.

Div. I—Prothorax scutiform, its lateral border visible from above and sharply
bordered.

Div. II.—Prothorax pulviniform (pillow-shaped), lateral borders in front
generally not visible from above.

Div. I was subdivided into five subgenera: Archarthropterus (= Sloane’s
wilsoni group), Arthropterus (= (?) Sloane’s parallelocerus group), Peltarthrop-
terus, Phymatopterus Westw. (= Sloane’s piceus group), Telarthropterus (= Sloane’s
macleayi group).

Div. Il was subdivided into three subgenera: Huarthropterus (= Sloane’s
denudatus group), Sticharthropterus, Panarthropterus (= Sloane’s hopei group).

He was apparently unaware of A. articularis Elst., described in 1919, while
much of his classification was founded on erroneous identifications. Thus he
considered A. macleayi (the type of Arthropterus W. S. Macl.) as identical with
parallelocerus Westw., these two species being taken as types of distinct groups
by Sloane, who writes: “There seems no objection to Kolbe’s two main divisions,
though his idea sometimes gives a false impression of relationship, e.g.,
A. denudatus has ‘scutiform’, A. kingi ‘pulviniform’ thorax. I think he knew
A. parallelocerus Westw., but was wrong in his identification of A. macleayi Don.
He does not describe his macleayi—Westwood says (Arcana Ent., ii, p. 8), ‘Mr.
Francillon’s unique specimen of this species is now in Mr. Macleay’s possession,
from whose figure in the work above quoted mine is copied’. The ‘work above
quoted’ is Annulosa of South Africa (W. S. Macleay), therefore Kolbe makes an
error when he says that Westwood’s figure was a copy from Donovan’s work. I
have seen the original type in the Macleay Collection and regard bisinuatus Macl.
as a synonym.’ Sloane notes the comparative dimensions of the prothorax of the
two species in question from the figures: A. macleayi Don. 6 x 6% mm.; A.
parallelocerus Westw. 7 x 8% mm. This prevents the acceptance of the name
Arthropterus for the other species placed here. My query as to the identity of

* An example (?) in Sloane Collection, Canberra, not described in Sloane MSS.

BY THE LATE T. G. SLOANE. 401

Sloane’s group with Arthropterus (Kolbe) is from Sloane’s uncertainty as to
Kolbe’s determination of parallelocerus with his own idea of that species.

Of Peltarthropterus and Arthropterus (Kolbe), Sloane writes: “I do not know
how to distinguish these from one another, chiefly because I cannot be sure of
any one of Kolbe’s species here, though am prepared to accept A. foveicollis as
right—I look upon Arthropterus Kolbe as equal to my parallelocerus group.”
Later (1/1/1928) is a note, “I cannot now consider his A. parallelocerus as equal
to the species I think it, by long ciliae of sides. He attributes no new species
here and I do not suppose that I know in nature one of the 3 species referred
here.”

Of Kolbe’s Division ii he writes: “I doubt whether there is anything more
than a mere analogical resemblance between the subgenera Sticharthropterus and
Panarthropterus in form of thorax.”

A. turneri Kolbe (néc Macl.)—‘‘not turneri Macl., seems likely to be the
common Victorian species westwoodi Macl.”

A. angulatus Kolbe (nec Macl.)—‘‘Macleay’s species is smaller than his and
is a Telarthropterus not a Peltarthropterus.”

A. humeralis Kolbe (nec Macl.) is given as a synonym of subcylindricus
Westw., without mention of Masters’ correction of the confusion with Macleay’s
species; but, according to Sloane, A. humeralis Macl. = nigricornis Macl.

Kolbe indicates four “gruppe” (Tijd. voor Ent., p. 12)—‘wyanamattae, turneri,
westwoodi and melbournei”’, on which Sloane comments: “Considering that in
nature one can hardly differentiate wyanamattae, westwoodi, and melbournei, it
seems impossible to accept these groups.”

A. waterhousei Macl.—‘‘placed by Kolbe in Huarthropterus belongs to
Panarthropterus.”’

A. mastersi Macl.—‘He is wrong about A. cylindricollis Macl. being the
do of A. mastersi: the type is a ? and is synonymous with mastersi.”

Kolbe also published a redundant name, A. dohrini, for A. howitti Westw.,
already replaced by howittensis Masters.

Of Kolbe’s species I find the following notes: A. secedens Kolbe.—‘‘For me
this is so near waterhousei Macl. that I cannot differentiate it by description.”

A. moretoni Kolbe—‘I do not think this differs from my westwoodi.”

A. ominosus Kolbe.—‘‘goes near westwoodi’.

A. sphinx Kolbe.—‘must greatly resemble, and be closely allied to, A. nigri-
cornis Macl. The prothorax in my specimen is longer than broad (2-5 x 2-3 mm.).
I hesitate to think they are different.”

A. ambitiosus Kolbe.—‘‘seems allied to rockhamptonensis Macl.”

A. daemelianus Kolbe.—‘‘very closely allied to rockhamptonensis Macl.”

A. geminus Kolbe.—“This seems as if it might be ? of A. Simson’s Bowen
species.”

A. pervicax Kolbe.—‘I identify for a Mackay species (Carter Coll.).”

A. suspectus Kolbe.—‘‘'Type with only one fore leg and no abdomen, I suspect
it of being ¢ of pervicaz.”

The above notes show the desirability of a revision of Kolbe’s species and of
their correlation with the others before any further work on the family is
undertaken.

The following new species described by Sloane are all of which I can find the
types. Others are referred to in the above notes, but these have either perished,
or are merged in other collections, at present unknown to me (H.J.C.).

402 AUSTRALIAN SPECIES OF THE FAMILY PAUSSIDAE,

ARTHROPTERUS OCCIDENTALIS Var. ORIENTALIS, N. var. (or Sp.).

Head convex, punctate; front tumid, neck wide, not contracted behind eyes,
antennae wide, basal joint transverse, inner angle prominent, 2nd unequal,
triangularly pointed on inner edge, 3-9 about 4 times as broad as long, apical
joint hardly longer than 8-9 together. Prothorax slightly broader than long
(2:1 x 2-3 mm.), subdepressed on disc, widest at anterior fourth, strongly and
roundly narrowed to apex, lightly narrowed to base, base angles obtuse, lateral
border well developed, stronger posteriorly, obsolete on anterior curve, punctures
strong and close, setae of pronotum very short. EHlytra parallel (6:7 x 3:6 mm.),
lateral row of fixed setae distinct. Dim.—11-11:5 x 3-6 mm.

Hab.—Vict.: Mulwala (Sloane); N.S.W.: Tullamore, Narromine, Warren,
Spring Ridge; Queensland: Roma (Fischer).

Resembles A. westwoodi Macl., but is at once distinguished by its fully
developed wings and much greater length of metasternum. I consider it the
eastern form of A. occidentalis Blkb., with the following differences: (1) Head
not at all depressed between eyes, (2) Prothorax not one-fourth longer than wide,
(3) Apex of anterior tibiae lightly emarginate—not triangularly excised.

ARTHROPTERUS LONGICOLLIS, Nl. SD.

é. Flightless, elongate. Head a shade wider than prothorax, punctate, front
convex, orbits not prominent, neck scarcely constricted behind eyes, antennae
wide, rather long, basal joint transverse, outer angle hardly prominent, 3-9 short,
about three times as broad as long, narrower than hopei, apical joint about two
and a half times the penultimate, evenly rounded in front. Prothorax (2 x 1:7
mm.) narrow, subparallel, widest at anterior fourth, arcuately narrowed to apex,
base wider than apex, lightly arcuate-truncate, setae very short and decumbent,
lateral border well developed and recurved. EHlytra (5:25 x 2:8 mm.) much wider
than prothorax, densely setigerous punctate, setae short and decumbent.
Metasternum rather long, transverse impression distinct. Legs wide, all femora
and tibiae more than twice as long as wide, anterior tibiae with front angles
obtusely pointed, post-femora slightly wider than tibiae, post-tibiae wide, exterior
angle produced backward in a truncate triangle, post-coxae cordate, apex obtuse.
Tarsi short, posterior with 2nd joint widely dilated, spongiose on inner side, 38rd
broader than long, a tuft of squamae on inner side, 4th small, broader than long,
dth longer than 2nd and 8rd together. Dim.—9 x 2-8 mm.

Hab.—New South Wales: Howell (Inverell district), from H. J. Carter.

Compared with type of A. elongatulus Macl., the size is larger and more robust,
antennae wider, less depressed above and below, more coarsely punctate, post-
tibiae not truncate at apex, but obliquely produced to outer angle. Allied to
A. macleayi Don., but differs by size larger, proportionally longer, punctures of
head, prothorax and elytra stronger. I do not think it is A. sphinx Kolbe, since
the posterior angles of prothorax are not ‘obtuse subrectis’, it is not setose
enough, also too small. Type in Collection of Division of Hconomic Entomology,
Canberra.

ARTHROPTERUS PLANICORNIS, Nl. Sp.

6. Winged. Reddish castaneous. Head (2 mm. wide) lightly strangulate
behind eyes, strongly punctate, punctures minute on neck, front hardly convex,
eyes large, not very prominent, antennae wide, basal joint quadrate, 3-4 succes-
sively widening, 5-10 equal, about three times as broad as long, apical broader than
long, parallel on sides, rounded at apex, about as long as 24 preceding joints, flat on

BY THE LATE 1T. G. SLOANE. 403

lower, convex on upper side. Prothorax subquadrate (1-8 x 2:15 mm.), widest before
middle, lightly narrowed to base, base wide, with truncate angles distant from pedicel
marked but obtuse, disc punctate, a little rugulose, setae minute on disc, few and
short on side. Hlytra greatly wider than prothorax (7 x 4 mm.), rather nitid,
punctures fine and rather dense, setae short and sparse. Legs of moderate width,
femora evenly arcuate on anterior side, post-tibiae hardly wider than femora, apex
oblique, outer angle obtuse, post-tarsi ordinary, joints 2 and 3 with spongiose
tissue beneath, 2nd large, but not twice as long as broad. Dim.—11 x 2 mm.

Hab.—New South Wales: Rydal (from H. J. Carter).

Allied to A. brevicollis Macl., from which it differs by antennae much wider,
apical joint equally rounded (in brevicollis the posterior side of this joint is
longer than the anterior, so that the apex is obliquely rounded). From A. howitti
Macel. it differs at once by the great reduction of setosity of antennae, prothorax
and elytra. It is the only species I know with the antennae widening towards
apex, described for A. subsulcatus Westw., but it has not “angulis posticis” (orbital
tubercles) “pone oculos valde porrecti’. Type in Collection of Division of
Economic Entomology, Canberra.

ARTHROPTERUS CONSTRICTICEPS, Nl. Sp.

g. Alate. Head wide, setigero-punctate, front convex, vertex subdepressed,
neck constricted behind eyes, antennae with joints 3-10 equal, less than twice
as broad as long, densely and finely setigero-punctate, apical joint subquadrate,
rather longer than broad, parallel on sides, lightly rounded at apex. Prothorar
subconvex, a little broader than head, widest at anterior third (2 x 2:5 mm.),
roundly ampliate on each side of neck, lightly and widely impressed near base,
disc a little depressed, median line well marked, lateral border narrowly reflexed,
obsolete towards apex and behind basal impression, punctures strong, separate
setae of moderate length, sloping backwards, sides rounded on anterior three-
fourths, lightly sinuate at basal fourth, base truncate. EHlytra parallel (7 x 4:2
mm.), shoulders rounded, a little prominent, punctation fine, setae short, sub-
recumbent. Metasternum large. Post-coxae elongate-cordate. Femora narrow,
not dilated at base, tibiae narrow, anterior more than twice as long as wide, apex
deeply emarginate, outer angle acute, post-tibiae narrower than femora, outer
angle triangular, rather prominent. Dim.—12 x 4:2 mm.

Hab.—Victoria: Mallee district (C. French, junior).

A single specimen in my collection, kindly given to me by Mr. F. P. Spry,
of Melbourne, ticketed ‘Mallee, 10-14’’.

A distinct species allied to A. latipennis Macl., from which it differs decidedly
by head less transverse, neck narrower and more evenly strangulate behind eyes,
antennae more slender, prothorax less quadrate, sides more rounded at widest
part, more sinuate to base, surface more setose. Looked at from the front the
prothorax is cupuliform, the anterior later slope of the sides continues on to the
neck, thus causing it to be constricted. As in latipennis, there is on each side of

the neck a deep transverse fossulet. Type in Collection of Division of Economic
Entomology, Canberra.

ARTHROPTERUS CRIBROSUS, N. Sp.

Head convex (2:6 mm. wide), vertex subdepressed between eyes, neck wide,
lightly transversely impressed, but not constricted behind eyes, punctures strong,
dense, extending on to neck. Antennae very wide, joints 3-9 about five times as
broad as long, 1 transverse, outer angle scarcely prominent, 3 unequal, inside

404 AUSTRALIAN SPECIES OF THE FAMILY PAUSSIDAE.

angle triangular, apical joint subhemispherical, seen from side rather thick,
upper side moderately convex. Prothorax short, much wider than head (2-4 x 3-2
mm.), widest before middle, rounded on sides, decidedly narrowed to base, basal
angles distant from peduncle, sides a little explanate towards basal angles, border
strongly reflexed, wider behind than in front, obsolete on apical curve, punctation
strong, dense, setae very short on pronotum, forming short, but strong bristles
on inflexed margins. Hlytra truncate, oval (7:3 x 4:3 mm.), punctures strong,
dense, lateral row present. Legs wide, post-trochanters cordate, post-femora very
wide, dilatate on inner side near base, upper side arcuate, the curvature bent at
basal third and sloping strongly to base. Anterior tibiae not twice as long as
wide, apex truncate on inner side, external angle triangular, not prominent, not
reaching as far forward as inner apical part, post-tibiae wide, short, setose-
punctate, outer angle wide, very obtuse. Metasternum short, very little longer
between coxae than length of post-coxae. Dim.—12°4 x 4:3 mm.

Hab.—N. Queensland: Kuranda (National Museum), Cooktown (Olive).

Flightless. Type in National Museum, Melbourne.

At my request, Mr. H. HE. Andrewes was good enough to compare this species
with A. foveipennis Blkb. and to send me the following note on it: “Belongs to
the westwoodi group and allied to westwoodi Macl., but the following differences
are apparent: form wider, antennae wider, basal joint with inside angle less
prominent, 3rd joint more sharply triangular on inner side, prothorax more
transverse, border wider and more reflexed towards base, basal angle obtuse,
not marked, elytra more densely punctate.’ As is usual in the genus, joints 3-10
of antennae of ¢ are more closely punctate towards sides, and have shorter setae.

Literature.
BLACKBURN, T.—Trans. Roy. Soc. S. Aust., xiv (1891), p. 68; ibid., xv (1892), p. 24.
BURMEISTER, M. H.—Mag. de Zool., 1841.
DESNEUX.—Wytsman’s Genera Insectorum, 1905.
Donovan, H.—Exotic Insects. Vol. iii. Epitome of the Natural History of Insects of
New Holland, New Zealand, New Guinea, Otaheite, etc. (London, 1805).
FAIRMAIRE, L.—Ann. Soc. Ent. France, 1xxii, 1905, p. 183.
Fow.Ler, W. W.—Paussidae of India. Fauna of British India, 1912.
GESTRO, R.—Ann. Mus. Civico Genova, xxxii (1892), p. 705.
LACORDAIRE, Th.—Histoire Naturelle des Insectes. Genera des Coleoptéres. Vol. ii
(1854).
LAMEERE, A.—Ann. Soc. Ent. Belg., xlvii, 1903, pp. 155-165.
Lea, A. M.—Proc. LInn. Soc. N.S.W., xxxi, 1906, p. 217.
.—Proc. Roy. Soc. Vict., xxiii, N.S., 1910, pp. 176-180.
Lerroy, H. M.—A Synopsis of the Classification of Insects (1913).
Mac.ueay, W.—Trans. Ent. Soc. N.S.W., ii, Pt. 2, 1871, p. 153; ibid., ii, Pt. 5, 1873, p. 337.
Mac.ueay, W. S8.—lIllustrations of the Annulosa of South Africa, p. 75 (London, 1838).
MULLER, R.—Ent. Zeit., xxiii, 1909.
RaAFFRAY, A.—Nouv. Arch. Mus. Hist. Nat. Paris, 2me Serie, viii, 1886, p. 307; ibid.,
2me Serie, ix, 1886, p. 1; ibid., 3me Serie, iv, 1892, p. 91.
SHaArpP, D., and F. Muir.—Trans. Hunt. Soc. Lond., 1912, p. 490.
WASMANN, E.—Notes Leyden Mus., xxv, 1904, 1-82.
.—Deut. Ent. Zeit., 1907, pp. 147-153; 561-566.
.—Tijd. voor Hnt., lv, 1912, pp. 257-260.
————..—-Zool. Anz., Ixviii, 1926, 25-30.
WeEstTwoop, J. O.—Hnt. Mag., v, 1838, p. 503.
————.—Trans. Linn. Soc. Lond., xviii, 1841, p. 581.
.—Arcana Entomologica, Vol. ii, 1845.
.—Proc. Linn. Soc. Lond., ii, No. 41, 1849, p. 55; ibid., ii, No. 44, 1850, p. 100.
Trans. Ent. Soc. Lond., N.S., ii, 1852, p. 84.
.—Thesaurus Entomologicus Oxoniensis, 1874.

USEFUL COCCINELLIDAE FOUND ON THE COMBOYNE PLATEAU.
By EH. C. CuisHotm, M.B., Ch.M.
(Hight Text-figures. )
[Read 27th September, 1933.1]

There are eight species of the family Coccinellidae, commonly called lady-bird
beetles, on the Comboyne Plateau, which are of decided economic importance,
seven of them being insectivorous and one a vegetarian, but nevertheless useful.
They appear first as the perfect insect from September to December, during
which time the whole cycle of the new generation is produced in most of them,
so that during these four months the eggs, larvae, pupae and imagines appear.
After December one seldom sees any of the species, except possibly Leis conformis,
and that in the adult stage. In a farming community such as this and in fruit-
growing areas, it is a very important matter to preserve and encourage any agent
which assists in destroying the many enemies of crops, whether cereals or fruit
or products of the flower garden, but unfortunately these small beetles are hardly
ever studied by the man on the land or, if they are noticed at all, they are looked
upon with grave suspicion as being identical with or closely allied to the Pumpkin
beetle (Aulacophora hilaris) of the family Chrysomelidae and killed on sight.

The species found are as follows: Leis conformis, Coccinella repanda, Verania
frenata, Verania (lineola?), Coelophora inequalis, Coelophora veranioides,
Catlineda testudinaria and Halyzia galbula.

Leis conformis (Text-fig. 1), measuring 6 mm., is of an orange colour, profusely
marked with black spots over dorsal surface of thorax and elytra. This is one of
the commonest species and generally appears first in September as an imago
feeding on several species of Aphis, viz.: Schizoneura lanigera (Woolly Aphis) of
the apple tree; the Aphis infesting the mint of the vegetable garden; Macrosiphum
‘rosae (Rose Aphis); Toxoptera aurantii (Orange Aphis); Aphis nerii (Aphis of
Foxglove); and a species of Psylla living on Acacia melanoxylon (Blackwood
Wattle) and probably other Acacias. This lady-bird is rarely seen after January,
having by then been through all its stages. The larva, if anything, is more
voracious than the perfect insect. The eggs are laid in clumps, are yellow in
colour and laid in an upright position on the leaves or stem.

Coccinella repanda (Text-fig. 2), measuring 5-5 mm., is of a general orange
colour with two black arrow-head markings on each elytron. This is a common
species here in the imago stage during the last three months of the year, at the
end of which time the second generation of imagines has made its appearance.
It is found feeding on Macrosiphum rosae on the rose, on Toxoptera aurantii on
the orange, on Aphis nerii on the foxglove, on Macrosiphum solanifolia on the
potato, on the Aphis on garden mint and the Psylla of Acacia melanozxylon. On
these plants it may be found in any-of its stages, the larva being very active as
an Aphis destroyer.

Verania frenata (Text-fig. 3)—This measures 5-4 mm. long, is of a general
orange colour with a broad black line where the elytra meet on the centre of the
dorsal surface and another broad black line running longitudinally along the

G

406 USEFUL COCCINELLIDAE FOUND ON THE COMBOYNE PLATEAU,

centre of each elytron with a right-angled projection inwards at the anterior end.
These markings are subject to considerable variation. Some specimens have no
internal projection of the black line on the centre of the elytron, and sometimes
the central line is connected at its posterior end with the broad line in the centre
of each elytron. This species is not often seen here. I have found it on foxglove
feeding on Aphis nerii and only in the month of November.

Verania (lineola ?) (Text-fig. 4) measures 4-5 mm., and is a general light flesh
colour with no conspicuous markings, except a central narrow dark line at the
contiguous edges of the elytra. This is a rare form here, only seen once on a
climbing rose bush, feeding on Macrosiphum rosae; this was in the month of
November.

Coelophora inequalis (Text-fig. 5) is 5:3 mm. in length. The general colour is
orange with four small unequal black spots on each elytron and these vary some-
what in size and shape in different specimens. It is a fairly common species found
in adult form feeding on the Aphis of rose plants (Macrosiphum rosae), of
foxglove (Aphis nerii), of mint, and on the Psylla of Acacia melanozylon; it is
found from September to November.

Coelophora veranioides (Text-fig. 6) is slightly smaller than C. inequalis,
being 5-0 mm. Its general colour is orange, with a central black line including
the inner edges of both elytra and a broad uneven broken line longitudinally
through the centre of each elytron parallel to the central line so that each broken
line appears as two stripes in the same line with an interval between them. This
is a rare form here, found as imagines only and in September and October,
feeding on Toxoptera aurantii of orange and Macrosiphum rosae of rose.

Callineda testudinaria (Text-fig. 7) is the largest of all the species under
consideration, measuring 6-8 mm. This is a fairly common form, seen first as
imago in September, and in all its stages during the succeeding two or three
months. It is extremely variable in its markings; of a general biscuit colour
with each elytron divided by narrow or wider black lines into five squares; some-
times the dividing lines are barely, if at all, visible, at others they are broad and
well defined. Its food consists of Macrosiphum rosae on the rose, Toxoptera
aurantii of the orange, Aphis neri of foxglove, the Aphis of mint, and the Psylla
of Acacia melanozylon.

Halyzia galbula (Text-fig. 8) measures 455 mm., is a bright canary-yellow
colour with two broad wavy black bands transversely across elytra, the anterior
one including the contiguous parts of thorax and upper edges of elytra, the second
posterior to it and passing transversely across elytra about their middle, also a
black spot covering tips of elytra in middle line at posterior end when these are
closed. This species, unlike all the others, feeds on the Oidium (Mildew) which
attacks the leaves of rose plants. This is a vegetable fungus which does great
harm to rose plants, especially the smaller-flowered climbing varieties. I have
also found it on grape vines, evidently feeding on a fungus attacking their leaves.
On one occasion in another district I found several on a grape vine which was
infested with the larvae of Phalaenoides glycine (Vine Moth), which were just
hatching from eggs, and very young forms. I have always suspected that they
were feeding on these larvae but it was only circumstantial evidence, for I did
not see the lady-birds actually eating the larvae, though they were on the same
leaves and the larvae had all disappeared in a few days.

If any of these species of beetles are in any numbers on an infested plant
it is a mistake to spray, as this method will certainly destroy the very agent

BY E. C. CHISHOLM. 407

which will clean up the pest in a much more thorough and efficient manner if
left alone.

Text-figures 1-8.

1. Leis conformis. 2. Coccinella repanda. 3a, 3b and 3c. Various forms

of Verania frenata. 4. Verania (lineola ?). 5. Coelophora inequalis.

6. Coelophora veranioides. 7. Callineda testudinaria. 8. Halyzia galbula.
xe2%

There are several beetles liable to be confused with these useful species. One
that should not be is the Pumpkin beetle, the only point of resemblance being the
black markings on an orange ground, but in every other particular, especially in
its shape, it differs considerably, being a long oval as against the rounded form
of the Coccinellid; also in consistency, the Pumpkin beetle being much softer,
especially the elytra, and more easily crushed. Another genus of the family
Chrysomelidae is Paropsis which very closely resembles the species of Coccinellidae
in general form but differs in having four-jointed tarsi while the latter has the
tarsi three-jointed. The members of the genus Paropsis are leaf-eaters. Among
the Coccinellidae themselves there are vegetable feeders, for example, some species
of the genus Hpilachna are vegetable feeders, some of them very injurious in the
vegetable and flower garden: EHpilachna 28-punctata feeds on the leaves of
Solanaceous plants including the potato, tomato, and other Solanaceae. This
species is about the same size as Leis conformis and marked in orange and black
in a very similar manner and may easily be mistaken for the latter, but Epilachna
is covered with a silky pubescence while Leis is smooth and shiny.

The dimensions given above are the actual measurements of specimens I had
in my possession, but they are subject in all to a wide range of variation. In
colour, too, they vary considerably. Those that I have spoken of as orange vary
from yellow through orange to almost bright red.

My thanks are due to Mr. E. H. Zeck, Assistant Government Entomologist,
and to Mr. H. J. Carter and also to the late A. M. Lea, Entomologist to the
Adelaide Museum, for identification of specimens.

MISCELLANEOUS NOTES ON AUSTRALIAN DIPTERA. I.

By G. H. Harpy, Walter and Hliza Hall Fellow in Hconomic Entomology,
Queensland University, Brisbane.

(Four Text-figures.)

{Read 25th October, 1933.]

Family LepripAr (Rhagionidae).
The name Leptidae is the one by which this family is known in Hngland
and Australia, but in America Rhagionidae has been adopted, following Kertesz.

DASYOMMA ABDOMINALIS, N. SD.

dg. Eyes contiguous, bare; the antennal triangle, face and occiput below are
light grey, the occiput above and the ocellar tubercle black. Antennae and palps
yellow with a few black hairs on the basal segments of the former, and dusky
hairs on the latter; elsewhere the hairs yellow. Thorax uniformly reddish-brown,
approaching chestnut colour but richer, and darkest on the pleura with all hairs
black. Abdomen uniformly shining black and with dark hairs. Legs dusky
reddish-brown.

Hab.—New South Wales: Como, ist October, 1921, found resting in a large
tunnel forming a water-channel that runs under the railway. The specimen is
unique.

Note.—Species of Dasyomma hitherto described from the Australian region
have the eyes hairy and are limited to Tasmania. Several species are now
known from the mainland of Australia, but only two are before me, both being
Gistinguished by the bare eyes and the colour pattern.

DASYOMMA FLAVA, N. SD.

9. Apex of proboscis, hairs on the palpi and on the two basal segments of
the abdomen and on the upper part of the occiput black; hairs elsewhere and
on the head yellow. Eyes bare. Thorax uniformly yellow with black hairs
above, the abdomen shining yellow and slightly dusky towards the apex. Legs
at trochanters and extreme base of the femora black or dusky, and the tibiae
black, but elsewhere yellow. Wings hyaline.

Hab.—New South Wales: Blackheath, 20th November, 1919, one female. I
have seen the male of this, but it is not before me. The large valley on the
west of Blackheath township has a creek near overhanging rocks from which
this specimen was taken; the rocks are adjacent to the footbridge crossing the
creek.

CHRYSOPILUS FASCIPENNIS, nN. Sp.
6. Eyes contiguous, bare, ocellar triangle black. Frons and face black with
a pulverulent covering that makes them shine dull yellow in accordance with
the incidence of light; the large bulbous tubercle in the centre of the otherwise

BY G. H. HARDY. 409

sunken face, and the head below are dull yellowish; occiput grey, and all hairs
white or yellow. Proboscis, palpi and antennae brown, and arista micro-
scopically pubescent. Thorax and abdomen black-grey with unevenly scattered
light golden tomentum and black hairs, whilst the pleura may have light grey
patches or even be entirely light grey, and all hairs there are white. Coxae
light grey with light golden hairs, otherwise the legs are yellow. The hyaline
wings have yellow-brown veins along which brown infuscations are conspicuous
at the apex of the two upper radial veins, the whole of the forked portion of the
two lower radial veins, at the apex of the discal cell and along the veins branching
from there, giving a general impression of an interrupted fascia across the wing-
tip, but sometimes these marks become joined at the base and sometimes they
are very faint.

9. Similar to the male, with well separated eyes and the head entirely light
grey or almost so.

Length, 10 mm.

Hab.—Queensland: Upper Brookfield, Brisbane; 7 ¢, 5 2. Mt. Glorious; 1 9.
December and January.

Note.——This, the largest known Australian Leptid, is common in the Queens-
land rain forests, and I have failed to trace it as a described species in the islands
north of Australia. The other Australian species have unmarked wings and are
small in size.

Family SrraATioMyYTIDAE.
Subfamily HrerMeEtIiIn AE.

There are six generic names attributed to species that occur in Australia,
and four certainly are valid and may be recognized by the following key:
IPAVVELUDESDINES HON) othe SCuUbeLMIMA yao sj eyevelaorel eve (elo /enciel oe belgnedein exavevolietede ois, «saraheusleve, dees «es 2

NLL OUtESPINESH-OnwebhexcSCuitellirnis spheres eeceoenciotetalcusloneke tichtnerei scheme ce Corea seal never avencnstes 4

2. With a pair of strong spines on thorax, one placed each side above wing insertions.
MEM Meat coe rcwenepan ean shee Reni kok ccaictaaecocomou static nen ape narteactiahavior st (easiest oral size) eres Negritomyia Bigot.

VWVat GL OUteESU Chia SDIMCS weer pecoctbnss taier: easy Rete chen crop cere rey cari auamevat cwe¥ons Louis, wystsheietone enero comic te 3
3. WVenation normal, M, being present ..... Elissoma White and Pycnothorax Kertesz.
Wed, Wi, BSG polopoodéacosdd bingo obo ope dgodu noone HHO DOO hOoouHodoDoDUGoUDOE gen.

AE cin alemwithian bLoad acute flaneel behinds themeyesme chore 4s. teenie
Pees Te May elaine cs eteviae Sone SIM aoa al ee sievaysar ey Lagenosoma Brauer and Perastomastix Enderlein.
TNEMMRNIES Ailsa Kooy Gy WENNER Geoodobocdb boon bouon coum doouEoo Hermetia Latreille.

I propose to use the first name whenever two names occur in the above key,
and to join the genus left unnamed under Elissoma, until adequate material
accumulates to show how further divisions are best formed.

Negritomyia and Hermetia are represented in Australia by one known species
each and the other two are plentifully represented. All genera have five fully-
developed segments in the abdomen, the others being more or less reduced and
retracted when at rest.

Genus Erissoma White.

White described this genus on a species that runs to other forms also
provided with scutellar spines, and Pycnothorar Kertesz belongs to the same
group and is probably congeneric. The form described below differs in having
vein M, missing, but it comes so close to White’s genus that it seems inadvisable
to separate it. Moreover, another species before me differs in having the
scutellum almost conical and raised to stand at right angles to the thorax, as
well as missing the vein M,. Indeed variations of this nature cannot be accepted
as of generic importance and it becomes very evident as more species accumulate,

410 MISCELLANEOUS NOTES ON AUSTRALIAN DIPTERA. i,

that the genera will need to be erected on better grounds than has been done in
the past..

ELISSOMA BRUNNEA, Nn. SD.

fg. On the antennae eight segments are traceable, but only six of them are
articulated and their lengths correspond to the following percentages: 11-2; 6;
26-6; 3-8; 11-4; and 41. The eyes are contiguous for about one-third of the frons
length. The abdomen has five visible segments of which the basal one is very
restricted and the widest part is at the apex of the third segment.

In colour the species is mainly brown, faintly varying to yellow-brown, but
white occurs on frons, on side of face and on anterior tarsi. Black occurs (or
the parts are considerably darkened) on the apical segment of the antennae, on
the ocellar tubercie, on the region behind the head, the neck and the small area
of the thorax adjacent on the humeral tubercle, on the pleura and posterior
coxae, trochanters and basal half of femora, on the first and two-thirds of the
second abdominal segment, the basal median area of the third and fourth segments
and the majority of the fifth segment. On the two paratypes the legs are entirely
yellow and the black of the abdomen not quite so extensive. The hyaline wings
have a black-brown stigmal area and a tinted central area. The third median
vein is missing.

Hab.—Queensland: Westwood, 12th December, 1924, 1 g, taken by A. Burns;
Ayr, 2 g, 24th December, 1930, taken by J. H. Buzacott.

Subfamily CLiTELLARIINI.
OPHIODESMA INNODUS Hardy.

This species was described from a pair, and since then four females were
taken from grass-trees adjacent to a small swamp north of Southport, Queensland,
in December, 1932, by Mr. L. Wassell. This brings the species into line with
O. flavipalpis, which also is only known to inhabit damp situations.

Subfamily PAcHYGASTERINAE.
Key to the genera of Pachygasterinae.

1 SCULEUIM. MWA ISPUM CS ee ees ye ark ores tur se (een leu ovis oe ce ces wana eer sucue heli oeMene tie waka neu eleven epomerec en irc Ueuteesaetan 2
Scmtelhume Swath oulySDINes sees yco scr: ce sreues ewes sci aire wous mente veureaeneiie eshene uel shots goueu sens fouemeae nel sree Mewevene 3
2. Abdomen of moderate length, not or hardly broader than thorax .... Hvaza Walker.

Abdomen very short and broad, almost circular, broader than thorax ............
eZ Ue oe sc ads Dee SRNR SRE EN Meee Sere oc fred e roy a erals Ait gc erst Sea SIO Or Ng Wallacea Doleschal.
Secutellum produced into a long finger-like process .......... Lonchegaster White.
SOOKE hbbaay savopmaoksl,) iaobayskeol’ Ene EWE ska omnocooduoucGobooUacoodbanduooKo bono & 4
4. Seutellum usually distinctly granulated along apical margin. Radio-median cross-
vein excessively short, practically eliminated. Third segment of antennae

(SU)

COHESperh evened nye fas] Ko) OND ULE Chet aes Sirs Wee epee SN ge acre re enero) ty Damaromyia Kertesz.
Scutellum not granulated. Radio-median crossvein long and normal. Third segment
of fantennae= much) longer tthan broad G2. es sec ces eieieieeie ens Dochmiocera Hardy.

Genus WA.LLAcEA Doleschal.

Of this genus I have seen two species, one being in the Ferguson collection.
The type of W. darwini Hill has not been examined by me, but the description
contains measurements that show the frons to be slightly more than one-quarter
the head-width on both sexes. In addition, the scutellum is provided with
about eight pairs of spines and a prominent disc-like process (prealar callus)
projects from the thorax. These disagree with the characters of the following
species.

WALLACEA SPLENDENS, 0. SD.

¢. Frons between 0-15 and 0:16 of the head-width and of normal shape,

having a transverse depression into which the carina runs. On each side of

BY G. H. HARDY. 411

the carina, out of this depression, there is a line of punctures running to each
side of the ocellar tubercle which extends to almost the whole width of the
parallel-sided frons. The long hairs of the tubercle are brown-black, and those
of the frons are similarly coloured but hardly discernible. A pair of silvery hair
spots at the base of the antennae. The black face has scattered dark hairs and
occasional silvery ones, together with a very thin white lateral border which
extends to the lowest corner of the eyes. Oral margin brown. Head, behind the
eyes, black with black-brown hairs and occasional silver ones. Antennae orange
with a white terminal arista which is brown at the base. Thorax thickly studded
with silvery-white hairs, these being depressed, whilst upright hairs are black-
brown. The silvery hairs, when seen from the front, form a thick mass extending
on to the scutellum, which has four small pairs and one large pair of marginal
spines arising from the scutellum and not from a specially formed flange as in
the species in the Ferguson collection. The silvery pubescence depends upon the
incidence of light to show its most vivid colour and at first may be found missing
from parts of the dorsum, but actually the hairs can be seen to form a dense
mass everywhere there except the lateral borders of the scutellum, where they
are entirely absent. The three basal segments of the abdomen are completely
covered on the upper surface with these silvery hairs, whilst they occur beyond
this in the usual pattern with the central part of the sclerite black and shining.
Other characters are quite normal to the genus.

®. The frons is between 0-26 and 0-27 of the head-width, thus corresponding
closely to that of W. darwini. There are more hair pits on the frons compared
with those of the male, and the white spots above the antennae are missing.
The number of silvery hairs is greatly reduced as in W. darwini.

The eyes, which have very scattered short hairs on both sexes, have a colour
pattern on the female (that of the male not noted) when alive. My notes and
sketch show a large red blotch covering the upper third of the otherwise green
area and this blotch has three parallel elongate marks, the middle one being
green like the main eye-colour, those above and below being blue.

Hab.—Queensland: Brisbane, 1931, 1 ¢ (holotype) in May, 1 @ (allotype) in
September, 1 2 (paratype) in November. I believe I am correct in allying the
sexes.

Genus LoNCHEGASTER White.

When in Melbourne last I examined and made notes and drawings of the
paratype of White’s species in the National Museum, but these are now mislaid.
Nevertheless I had concluded at the time that the two Queensland species before
me were not conspecific with White’s, although one was very close. As far as I
can now judge, the only specific character that will readily distinguish the three
forms lies in the scuteilum, which is somewhat triangular, with a terminal
elongation which slightly converges towards the apex, where it is broadly
rounded. One of the Queensland forms has the scutellum strongly raised, much
like that of the typical species, whilst the other has it lying in a plane with the
thorax.

LONCHEGASTER DECUMBENS, 0. Sp.

9. Black with yellow antennae, legs and wing veins. Frons converging
slightly towards the antennae, broad, a little less than 0-29 of the head-width and
with uniformly scattered hairs each side of the slender carina that tapers to the
part where the frons recedes near the antennae and where two silvery-yellow

412 MISCELLANEOUS NOTES ON AUSTRALIAN DIPTERA. 1,

hair spots lie. These hair spots extend down each side of the strongly receding
face as a rather broad stripe. Behind the eyes the head is strongly swollen,
giving a broad posterior orbital margin. Thorax rather densely covered with
hair-pits from which arise strongly depressed yellowish hairs forming conspicuous
pubescence over the dorsum. The hairs of the pleura are scanty and black. The
scutellum lies practically in the same plane as the thorax and is studded with
hair-pits and black hairs. It is triangular in shape, being as long as the width
at the base, from whence, slightly on the dorsal side, it is produced, finger-like,
reaching about half as far again. This prolongation is nearly three times as
long as its maximum width. The abdomen is more brownish than black, and
probably abnormal, but it has less dense hair-pits and black-brown hairs.

Hab.—Queensland: Brisbane, October, 1924, 1 9.

Note.—Another specimen before me is identical, but the scutellum is raised
and so presumably it belongs to another species. It is a female from Brisbane,
October, 1923.

DocHMIOCERA AUREOLINEATA Hardy.

6. Like the female but differs in having the eyes contiguous along the
central third of the distance between the ocellar tubercle and the insertions of
the antennae. Also the antennae differ in having the arista placed terminally.

Hab.—Queensland: Bunya Mt., 1 ¢ (allotype), 3rd January, 1926, taken by
Dr. A. J. Turner. There are a number of females in collections, but this is the
only male I have seen.

Family TABANIDAE.
Genus PELECORRHYNCHUS Macquart.
A good deal of unpublished work has been done on this genus by the late
E. W. Ferguson, by Dr. Mackerras, and by myself, with a view to combining it
in a complete revision describing many new species. The following key forms
one of my contributions to that abandoned effort:

il, ANlokoseaveral, Oye Wave GOS Shamileie shay royoyboe GoRwWA oooaoduacbuvcddouuood0doucuo0UE 2
Abdomen of the sexes differing in colour pattern, that of the female being orange
With nthree DIACK Str ies ee eee co Bos cue ee serra eee Ne a aie eee onal ee ouney oere Group 3

WH, AN oeKoveavsror Joysyavokercl Ayraliloy loehoe len: Goo5uugoccbooodboduucodguooude Group) 13
Abdomen uniformly shining black; bare or practically so ............. Group 2

Group 1.

3. Abdomen with hair bands alternating with complete white bands .............. 4
Abdomen with hair bands alternating with interrupted white bands, or the white
almost) seliminatcedis a micapoeieucesea we te es) ailcr Seals acie ieoneeeee Rec TR See. ito rore, Crea Oona to ire 5

4. Spots on wing large and confluent so that only six are usually separated. With
some red hair on cheeks and on the anterior part of the pleura ..............

REIS Sits OL CREOLE CEO RCRA OLDEST MCU ATO Oho EGG met orrssainies Choe nigripennis Ricardo.

Spots on wing smaller, few being confluent, so that ten or more are separated. All
hairs) ofmeheeksvandpleunawy.ellows . oes aici s ie eieie ee personatus Walker.

5. Abdomen with white bands interrupted in centre, dividing the white area into two
BLES Ee Re SUP LMR S, aes tre] etiaiee watranis.retcloas sheet eee Se PS Wee eas NE Heer Mtn ST haath AUR De) ae Ne eRe eRe 6
Abdomen with white bands interrupted each side of centre, dividing the white area
into) three) parts) or the awhite practically, veliminated 2544 eee 9

6. A white spot confluent with a white stripe on the black stripe of thorax. Wings
spotted: JAbdomen -withoultered® hairs —.o.5.¢ 0 oe one ee albolineatus Hardy.

At most with only a white spot on the black stripe of thorax ................ 7

(a Abdomentwith) red shairs ss iWwinesnspotted: A sanction eristaloides Walker.
Abdomen swithoutsred hairsy wines not Spottediers imc seis soe ore eee cena 8

8. With conspicuous white spot on black stripe of thorax. No black median mark
AdjacGenticto: (SGucell wre See ee areas tee oicetcicoude ete ate eel ae arcarecep eens montanus Hardy.

With spot on black stripe of thorax not white. A black median mark adjacent
Reaxets(o) bis o1 Uhh calwMetdna cicaemsseto tab or Gia oro ONCE re kaa eile ce eerie ota oO widiceckdots occidens, Nn. sp.

BY G. H. HARDY. 413

9. Interrupted white bands conspicuous and well defined. Hair bands with black

HAIG SE Re COMM AUT Camecicincradcueteiier se sushi ierot bacnetere trate meie ene eveueroetenenehele olivei, n. sp.
The white on abdominal bands reduced to inconspicuous and ill-defined spots. Hair
bands with fiery red hairs predominating .................- igniculus Hardy.

It is not proposed here to review the second and third groups, for these have
not come in my special studies and new forms need to be incorporated within
one of them. Those already described fall.into the following:

Group 2: fusconiger Walker (that of Taylor from Queensland is not
conspecific) ; claripennis Ricardo, tillyardi Taylor, dequeti Hardy, and flavipennis
Ferguson.

Group 8: fulvus Ricardo, distinctus Taylor, and mirabilis Taylor. The first
two have the abdomen on the male entirely black, and the third with alternating
hair bands and white bands, the latter being broadly interrupted in centre and
at sides, restricting the white to two rather small spots on each band.

Group 1 is well represented in Tasmania, much more so than on the mainland;
only one species of group 2 is known from Tasmania, and none of group 3. The
full list is as follows:

P. nigripennis Ricardo, Cradle Mt., Jan., 1925 (A. J. Turner), (also Victoria

and New South Wales)..

albolineatus Hardy, Cradle Mt.

eristaloides Walker, Strahan, Jan., 1924, a long series of females found
ovipositing in the perpetually damp mud of the track in People’s
Park; Zeehan, one male on a swamp and probably a stray specimen;
Huon District, Dec. and Jan., 1914 and 1916, two males.

montanus Hardy, Mt. Wellington, Jan., 1916 and 1924.

occidens, n. sp. Zeehan, Jan., 1924, and Cradle Mt., Jan., 1917.

olivei, n. sp. Strahan, Jan., 1924.

igniculus Hardy, Cradle Mt., Jan., 1917, and Zeehan, Jan., 1924.

fusconiger Walker, Wynyard, Jan., 1915, and Ulverston.

PELECORRHYNOCHUS OLIVEI, N. Sp.

A small species with spotted wings and three white spots on the second to
fourth abdominal segments.

6. Eyes contiguous. Face with black hairs blending into the yellow beard.
Palpi with black hairs. Antennae black with the basal segments reddish. All
hairs behind head yellow. Dorsally the thorax is deep brown covered with mixed
black and yellowish hairs. There are two white, well separated stripes lying
practically along the whole length, and a short triangular central stripe reaching
the scutellum and lying along the apical quarter. The scutellum has a small
white area, but the majority is covered with a broad brown border, uniformly
wide, and is clothed with long black hairs, except below, where they are yellow.
Some yellow hairs occur at and above the wing insertions, and some white
hairs on the postalar tubercle, these blending into the white pleural hairs which
are so abundant and long that the ground colour is not perceptible.

The abdomen is entirely deep brown on the first segment except for the
whitish but obscure lateral spots. The dorsal area of the three following
segments is brown with three large white spots on each, whilst ventrally they
are white with a narrow black apical band. White hairs occur on the white
areas and black hairs on the brown and black areas, but the apical margins of
the segments are bordered with bright yellow hairs. The wing spots are smaller,
but similar to those on P. personatus.

414 MISCELLANEOUS NOTES ON AUSTRALIAN DIPTERA. 1,

Hab.—Tasmania: Strahan, February, 1924, a long series from two small
swamps adjacent to ‘“‘The People’s Park’. The type series is a very long one,
of which twelve are still before me.

Note.—It would seem that the various species of Pelecorrhynchus that occur
in Tasmania are mainly restricted to swamps of certain types. No other species
was taken on the same swamps as these, nor were they seen away from these
small areas of a few acres each. This species is named in honour of Olive
Hardy, my wife.

PELECORRHYNCHUS OCCIDENS, Nl. Sp.

P. montanus var. a, Hardy, Rec. Austr. Mus., xiii, 1920, 34, fig. 3.

Very similar to P. montanus but quite distinct in markings. A diagram of
the differences was given in my earlier paper, where it will be noted that the
dorsal stripes of the thorax are much wider apart, less irregular in outline and
the spots that interrupt them at the suture are less intense. Also there is a
remnant of a central stripe adjacent to the scutellum. The lateral and median
bands which are joined at the shoulders in my figure are incorrectly given, but
there is a tendency towards the character. I have found no other differences
between the two species.

P. montanus is only known from Mt. Wellington, occurring on the large
swamp at the summit, whereas the present species was first recorded from Cradle
Mt., and is now known to extend westward to Zeehan. The species inhabits the
fringes of large swamps and is taken resting on or flying over the foliage of the
taller shrubs, whereas the species associated on the same areas are distributed
over the swamp.

Hab.—Tasmania: Cradle Mt., January, 1917, where it occurred in abundance,
and Zeehan, January, 1924, a small series, of which 2 ¢, 2 2 are now before me,
the others being distributed in various collections. These were taken at the
large Swamp adjacent to the township and were somewhat scarce and hard to
catch. The holotype and allotype are those in the Australian Museum, Sydney,
and originally labelled “montanus var. a’.

Family BomMByLiiDAk.

With the exception of the Lomatiinae, this family was revised by Mr. F. H. S.
Roberts and, partly due to his efforts, there has been some advance in the generic
relationship of the genera Comptosia and allies. The species placed under
Oncodocera in Australia do not belong there but apparently will form a new genus.
The species retained under Comptosia form a bewildering array that seem to form
a number of complexes, one of which was combined as a single species by me
under the name Comptosia sylvanus Fabr. That name, as used, can have no
standing, for it is uncertain if Fabricius has the genus, for it does not appear
to occur during the period when Banks collected, but makes its first appearance
about a month later. Three species were definitely combined under the name.

Comptosia hermeteles Schiner is the only one found in the Brisbane area
and extends as far south as Sydney, where it is the first of the genus to appear
on the wing.

Comptosia geometrica Macquart, from Tasmania, would appear to be the
same species as the common form in Sydney and does not reach Queensland.

Comptosia tricellata, Macquart’s second species from Tasmania, would seem
to be limited to Mt. Wellington.

BY G. H. HARDY. 415

Complications start when an attempt is made to bring the abundance of
material from the mainland into conformity with the two Tasmanian species. To
all appearances they become mingled both in characters and in distribution.
Field notes made on the various forms in Australia might aid in the disentangle-
ment of this complex.

Comptosia lateralis Newman is the male of Comptosia ducens Walker, founded
on the female. There are several other known cases where the white fascia of
the wing is limited to one sex, cases being frequently taken in copula by Mr.
F. A. Perkins at Stanthorpe, Queensland, and by others.

Genus Puruirta Meigen.

Roberts has drawn attention to P. hilaris Walker as being a very variable
species, and he believes some of the females standing under the name may belong
to other species, as yet undefined. Probably there is a complex here, for I can
separate, on male characters, two species, of which the females might be readily
mistaken for Walker’s form. The following key will readily separate these on
the males.

1. Proboscis elongate, about three times the length of the head. Face and frons with

white hairs on the male. Western Australian species .... albocapitis Roberts.
Proboscis normal, about twice the length of the head. Eastern Australian species 2
Zmeacesand trons with) black hairs on) the male 2. 3s.2...6..-s6002455.6 hilavris Walker.
MaCewandeshrOnstawitheivibitemh ainSeon eth esemaley iyo otelarctencienehol clever encore) lets ele) eter leet 3

3. Yellow or almost entirely yellow species, even the black of the thorax being strongly
UMN Se Gewese yie LOWE faces ak veseu ene he cele, rane feoudey cuer oman ocaien aera ledepone ase er sieueuay Ss flava, n. sp.
Mostlyvablackiawathipy.ellow, srmalrlain SSigeWencencisicqcten sens ein si citiay« eveienenel eel ticere nigrina, N. Sp.

I do not see any means of separating the females of the three species before
me, except in the case of P. flava, where the two basal segments of the antennae
are yellow, while in the others they are black. It will need some very careful
field work and, when possible, collecting species in copula, before more of this
complex can be satisfactorily solved.

PHTHIRIA FLAVA, N. Sp.

6. Head yellow and entirely with white hairs. Eyes contiguous. Antennae
with only the third segment black. Proboscis black and about twice ‘the length
of the head. Palpi normal, black, reaching to about half the length of the
proboscis. Thorax black-grey, strongly tinged with yellow and bearing sparse
yellow hairs. From the humeral tubercle to the postalar callus, and the region
near the scutellum, the whole of the scutellum and a small part of the pleura
are yellow. The abdomen may be entirely yellow or black at the base, with one
or two small median spots beyond this. Legs, except coxae, which are black-
grey, and the halteres and the veins of the wings are entirely yellow.

©. Similar to male, but the eyes are widely separated and the yellow of
the thorax is much more extensive, the pleura and coxae being almost entirely
yellow. The abdomen entirely yellow. The black region behind the eyes fades
into the broad yellow margin behind the eyes as on other species, but the black
seems much more restricted.

Hab.—Victoria: Ouyen, January, 1931. 7 3g, 1 9. This species was very
abundant in one spot in the Mallee country, but time did not permit observation
of it.

PHTHIRIA NIGRINA, N. SD.
do. Like the preceding species, except that the antennae, the thorax including
the scutellum, most of the femora and most of the abdomen are black. The hairs

416 MISCELLANEOUS NOTES ON AUSTRALIAN DIPTERA. 1,

of the head and pleura are white, elsewhere yellow and no black hairs are to be
Getected.

9. Similar to that of the preceding species except that the antennae are
entirely black and the yellow of the thorax is mueh less extensive, being reduced
to two spots just above the yellow scutellum, the postalar callus and a small
adjacent area extending on to the pleura. The coxae and most of the legs are
yellow, but the hind tibiae and tarsi are black. On both sexes the segments of
the abdomen are black but heavily margined with yellow.

Hab.—Victoria: Ouyen, one pair in copula, which forms the holotype and
allotype, and three male and one female paratypes, January, 1931. This species
is very abundant in the Mallee, but I mistook it for P. hilaris and did not collect
many.

Family APprmoceRIDAE.
Genus AprocERA Westwood.

There are six specific names attributed to species in this genus, which is a
very large one in Australia. The typical form A. fuscicollis Westwood, 1835, may
be regarded as being conspecific with Laphria brevicornis Wiedemann, 1830, and
attributed to the common form occurring around Sydney. Although Pomacera
bigotii Macquart, 1847, is attributed to Tasmania, that is possibly an error, as the
genus is not known from there. The illustration does not agree with any form
I have seen.

A. maerens Westwood, 1841, is readily recognized, as the palps and proboscis
were illustrated by Westwood, and there is only one species in collections that
‘agrees with it.

All the above belong to the sturdily built forms, and one of the more slender
types, another Sydney species, has been attributed by somebody (? Froggatt) to
A. asilica Westwood, 1835.

Hermann’s species A. vulpes, 1909, is not known to me but should be recog-
nizable by its description. ie
: Williston (1908) illustrated a wing of a species which he regarded as being
a new genus from Australia but did not give it a name. This species, for which
I have searched for many years, proves to be very variable in venation and in
no way distinct generically; it is named and described below.

APIOCERA MARITIMA, Nn. Sp. Text-fig. 1.

This fiy has the thick pulverulent covering, whitish in colour, that makes
the ground colour hardly discernible and there is no pattern. The basal segments
of the antennae, the palpi and the legs are tinted with brown, but the third
segment of the antennae, the eyes and proboscis are black. The hairs throughout
the head, body and legs are light grey. Normally on the female the seventh
segment of the abdomen is retracted within the sixth, and it is brown, like the
eighth, which protrudes beyond the sixth. The prosternum is small and is
surrounded by a membraneous area that is covered with hairs and hence has the
appearance of being a sclerite and not membraneous in texture. The venation is
more or less, and sometimes considerably, reduced, but it varies towards the
normal type; any of the veins, the fourth and fifth radian and the median, may
be short or even missing. The diagram given here to show the character of the
head appendages will enable the fiy to be reliably determined even if other
characters be ignored.

BY G. H. HARDY. 417

Terminaliia.—There are two well-defined lamellae on the ovipositor and these
are placed in the normal position, being contiguous along the median line.
There is a structure that seems to represent a hinged median plate, the two
together being situated at the base of the anal papilla, forming a V-shaped depres-
sion there, but this is not apparent until the anal papilla is exserted and the two
plates, forming the arms of the “V”, are drawn apart. The gonopore is situated
within a genital groove that is bordered by a chitinous ridge that surrounds it
on three sides. This ridge represents two lateral arms of chitin joined by a
cross-piece at the base to form a single sclerite. The ninth tergite has at least
eight spines placed along the apical edge each side of a strong upstanding dorsal
ridge. The eighth sternite appears to be simple, but part of it is turned inwards
so that the cleft at its apex becomes hidden within the genital cavity.

On the male, both upper and lower forceps are present, the claspers being
fused to the latter. The lamellae are formed by three long plates, the upper ones
narrow, the lower one being very broad, translucent and dark at the tip. The

Text-figs. 1-4.
1. Apiocera maritima, n. sp. Lateral view of the head.
Scenopinus civiculus, n. sp. The venation, the head seen dorsally and laterally, and
the antenna, seen dorsally, of the female.
3. Pseudatrichia mariaensis, n. sp. The venation, the head of both sexes seen dorsally
and laterally, and the antenna.
4. Paraliptus mirabilis Bezzi. Lower portion of the head showing the flange that
borders the eye below.

i)

418 MISCELLANEOUS NOTES ON AUSTRALIAN DIPTERA. 1,

other characters were not made out with assurance, but the aedeagus seems to be
short.

Hab.—Queensland: Southport, November and December, 1931, 5 J, 3 9, all
taken on the coastal dune along the foreshore of Ocean Beach, Burleigh, 9th
January, 1932; 1 2 in the collection of Mr. B. Blumberg.

Note.—This is one of the sand-coloured flies that frequent sand-dunes and all
of which have the same light grey pulverulent covering, often tinged with brown.
With it occurred a similarly coloured robberfly of the tribe Clinopogonini and a
smaller one of the same tribe not so coloured, also Anabarrhynchus maritima
Hardy (Therevidae) and Sarcophaga litoralis J. and T., besides a few Acalyptrate
flies. Later, when Apiocera maritima became scarce in December, males of another
became plentiful, a rather black species, of which I have not seen the female.

Family ScENOPINIDAE.
The three species of this family that I have found in Australia belong to two
genera, recognizable as follows:

Wings with the vein R, running free to the wing margin ............ Scenopinus Latr.
Wings with the first median vein meeting the vein R, before it reaches the wing
SO OVE =3) GUN mera ere TAD AA ea SL gy GeO ANG ATE RU at RG PP RC gues a ic ene Pseudatrichia O.-S.~

Scenopinus Latreille.

I believe S. fenestrata Lin. has been reported from Australia, but I have
failed to find the reference and it is not in any Australian collection I have seen.
From New Guinea Omphale papuana Krober is described and also Scenopinus
birot Kertesz. In the Hawaiian Islands, Greenshaw recorded Scenopinus nigra
Degeer, a species of the Northern Hemisphere. I have been unable to ally the
following species to these.

SCENOPINUS CIVICULUS, nh. sp. Text-fig. 2.

9°. Head as broad as thorax, black. Frons not quite one-third the head-
width, mostly shining and smooth, but slight punctures occur in two rows,
irregular in shape, each side of the median depressed line. The punctures extend
over the ocellar tubercle and some slight wrinkles occur, mainly on the lower
half of the frons. Behind the eyes there is a well developed flange, shining and
with but few punctures, these being mainly confined to the lower half. Seen in
profile the frons extends a little in front of the eyes, being broadest at the ocelli
and disappears at the antennae. The oral cavity is large, leaving but little of the
face to be seen, this being covered with a slightly grey tomentum. The hairs on
the cheeks and elsewhere are somewhat yellowish. Antennae brown, with the
third segment long, and a marked depression near the apex on the outer side.
The eyes, in life, have a straight thin transverse colour band, placed so as to
point slightly above the antennae. The band reaches across the eye from margin
to margin.

The thorax is black, fairly densely covered with punctures and wrinkles, the
slight hairs are somewhat yellowish. The scutellum is semicircular, similar to
the thorax in other respects and with a subapical depression bordering the outer
margin at about two-thirds its length. Below the scutellum and on the post-
humeral callus, the colour is yellowish-brown.

The shining black abdomen is sparsely punctured, somewhat depressed, and
about as long as the head and thorax united; it is conspicuously broader towards
the apex. Legs brown, apex of the tarsi stained with black. The hyaline wings
have yellowish-brown veins and the halteres have a white club. Length 5-6 mm.

BY G. H. HARDY. 419

Hab.—Queensland: Brisbane, November to March, 1924, and subsequent year's,
5 2 in my collection and 1 ? in that of Mr. F. A. Perkins. New South Wales:
Sydney, 26th November, 1920, 1 9. South Australia: Adelaide, January, 1931, 1 9,
and another without data. Mostly from windows.

PSEUDATRICHIA Osten-Sacken.

This American genus was not recorded from any other part of the world till
1922, when I recorded it as Australian. There are two species taken by me, one
being from Sydney, probably La Perouse, on Ist December, 1918, but this small
brittle pair taken in copula has completely broken up, the head, thorax, wings
and legs of the male being all that is left. It was about half the size of the one
described below.

PSEUDATRICHIA MARIAENSIS, n. Sp. Text-fig. 3.

9. Head as wide as or slightly wider than thorax; elongate, so that seen
laterally it is somewhat elliptical. Frons wide, one-quarter the width of the head
at its narrowest part. The head behind the eyes is very bulging, no flange is
formed and so the cavity behind the eyes is slight. Seen laterally the ocellar
tubercle stands above the eyes and the frons is mainly hidden. The frons is
shining, but there are two rows of punctures each side of the median depression;
these punctures stop before reaching the ocellar tubercle. In conformity with the
elongate head, the face recedes abruptly. The hairs of the head are more abundant
than those of the preceding species and somewhat yellowish. The antennae have
the third segment only two and a half times the length of the first and second
segments combined, rather conical in shape and with a small depression at the
apex.

The thorax is black, rather densely covered with punctures and yellowish
hairs. The semicircular scutellum is scarcely punctured, shining and bulging,
being divided from the thorax by a deep depression. The posthumeral callus has
a strong tendency towards being brown. The black abdomen is conspicuously
wrinkled, considerably longer than the head and thorax united, and widest at
nearly half its length. It has a vestiture of rather long, black, moderately
abundant hairs towards the apex. The legs are black, the wings hyaline, with
brown veins. The halteres are black.

g. Agrees with the female in general shape and colour, except that the
abdomen is metallic blue-black with white edges at apex of segments. The frons
is reduced in the centre so that the eyes are nearly approximate there. The post
orbital margins are prominent.

Length of both sexes, 5 mm.

Hab.—Tasmania: Maria Island, 29th December, 1915, 1 g, 1 9, in perfect
condition, taken in copula, probably by sweeping along the shore or close to it.

Family DoLicHOPoDIDAE.
PARALIPTUS MIRABILIS Bezzi. Text-fig. 4.

The type, in the Australian Museum, does not conform to Bezzi’s illustration,
which is too attenuated, and in addition there is a flange, bordering the lower
portion of each eye at the rear, that Bezzi mistook for palpi. I have given here
a camera lucida drawing of the region from the lateral aspect and also from
another angle which shows the flange is detached from the facial region. The
palpi themselves are hidden in the oral opening, which is choked with sand.

420 MISCELLANEOUS NOTES ON AUSTRALIAN DIPTERA. i.

On the type, the biserial acrostichal bristles occur as far as the insertion of
the pin, where the area may or may not be flat. According to my key (1930),
it runs to Dolichopus or to couplet 14, in accordance with whichever the doubtful
character may be. The anterior legs are raptorial. I was unable to place the
genus in its subfamily, but Bezzi considered it to be Hydrophorinae.

Family SyRPHIDAE.
CERIOIDES ORNATUS Saunders.

This species is associated with bee-hives. Mr. H. Hacker took a series of
females at the entrance of hives and I once met with it in this capacity by two
males hovering over a hive, and the bees were in an agitated condition until the
flies were caught. Evidently it was the humming sound that upset them. More
recently Mr. L. Wassell bred it from larvae out of waste brood comb. Half the
flies to emerge did so normally, but the remainder failed to form their wings
and the scape did not develop, the antennae remaining attached in appearance
at the head level, and not in advance of it. Mr. R. Veitch has drawn my attention
to a pair in the Department of Agriculture, Brisbane, each being labelled “one of
five at nest of Trigona carbonaria, 16.2.1927, H. T(ryon).” There can be little
doubt that it lives in association with native honey-gathering bees.

CERIOIDES SUBARMATUS C. and B.

This occurs in Brisbane in the very early spring (August), and can be
found at Leptospermum when the flowering season is not delayed. At the flowers
it looks very like a wasp, not only in its general shape, but also on account of
its habit of folding the wings longitudinally in the manner of the Diploptera. At
this time of the year the wasps are absent or but very rare, for it is too early in
the season for them.

XYLOTA IRRIDESCENS Ferguson.
3. Like the female, but the eyes are contiguous for a short distance.
Hab.—Queensland: Brisbane, 1 ¢, the allotype, August, 1927, at Leptosperm.
The species was originally described from a unique female. There are no others
known. The fly, on the wing, looked so like Lissopimpla semipunctata Kirby, that
I at first mistook it for that common ichneumon.

THE GENUS PTEROSTYLIS R.Br. [ORCHIDACEAE].

A NEW SCHEME OF CLASSIFICATION, WITH NOTES ON THE DISTRIBUTION
OF THE AUSTRALIAN SPECIES.

By the Rev. H. M. R. Rupp, B.A.
(Thirty-eight Text-figures.)
[Read 25th October, 19338.

Difficulties of determination in the genus Pterostylis arise largely through
the fact that affinities and points of resemblance between the different species
are curiously complicated, features of association between certain forms being
accompanied by divergent features which suggest quite other relationships. As
a result it is probable that whatever characteristics be taken as the bases of
the primary and secondary groups of the genus, opinions will differ as to whether
other characteristics would not have served the purpose better. The aim of the
classification proposed in this paper is simply to provide a clearer and more
direct route to the identification of species than is now available: and whatever
criticism may be fairly applied to it, I think it will be conceded that here a
strange form may be readily traced from larger to smaller groups until it is
either isolated, or restricted to the company of a few other species. At that
point the scheme ends, and ultimate descriptions must be sought in existing
publications. Accompanying the table of distribution of Australian species which
follows the scheme, I have given references for each species; and in selecting
these references I have been guided by practical considerations, believing that
a reliable illustration of the plant or its flower is a very great help towards
determining identity.

A few non-Australian species of Pterostylis are unknown to me, but I have
in my herbarium most of the New Zealand forms which have been described,
and some which are still awaiting description. All these can! be fitted into
the new classification, and from such information as I have been able to glean
I think the scheme provides a place for all known forms.

Robert Brown primarily divided the species known to him according to
the character of the appendage found at the base of the labellum. This basis
of classification seems more appropriate for use in subdivisions than in primary
sections, but in any case it presents difficulties which have increased with the
number of species. In Bentham’s “Flora Australiensis” we are given two primary
sections: (1) Antennaea, in which the lower sepals are erect with their free
portions embracing the galea, and (2) Catochilus, in which they are reflexed
in front of the ovary. This is a very important distinction, especially if we take
“reflexed” as equivalent to ‘deflexed or spreading’; but it involves certain
anomalies in the subdivisions, and in particular is open to the following criticisms:
(1) The classification includes in Catochilus two species (P. barbata Lindl. and
P. turfosa Endl.) which have hardly anything in common (beyond generic

H

422 THE GENUS PTEROSTYLIS,

character) with their associates, except the attitude of the lower sepals, and
which indeed in the character of the labellum and other features differ strikingly
from all other species in the genus. (2) In both Antennaea and Catochilus we
find species with (a) basal rosette or cluster and no stem leaves, (0) stem leaves
and no clearly recognized basal rosette, (c) basal rosette (or cluster) plus stem
leaves. This duplication is perplexing to those not well versed in, the genus,
since these features, discernible at a glance, at once suggest a means of narrowing
the search for a species down to one of three groups, yet the groups are found
to be split up between the two sections.

In the scheme set out below, it may be objected that Catochilus itself has been
split up, not only between the two primary sections, but also among the
subdivisions of Section A. This is true, yet this distribution will be found
less confusing than the anomalies referred to above; and it will be found
that the new scheme permits the segregation of the most closely allied forms
of Catochilus into groups just as before. The feature upon which Bentham’s
sections are based is too important to be lost sight of; I have retained his terms
Antennaea and Catochilus, which will be found bracketed against the groups
associated with them.

I had decided to omit from the Australian species named in the scheme
P. clavigera Fitzg., P. pyramidalis Lindl., and P. Suttonii. In deference, however,
to the opinion of Dr. R. S. Rogers I retain the first two, at the same time giving
my reasons for my former intention. P. clavigera does not appear to have been
seen by anyone since Mr. A. G. Hamilton collected it near Mudgee more than
40 years ago. Mr. Hamilton took it for P. nana, and although no specimens are
known to exist, drawings and descriptions are at least hardly adverse to this
opinion. Dr. Rogers, however, has reminded me of Fitzgerald’s unpublished
plate in the Mitchell Library at Sydney, and it must be admitted that the details
—particularly of the column—are not reconcilable with the type of P. nana.
I agree therefore that until fresh material is available we should give the species
the benefit of such doubt as may exist about it. The case of P. pyramidalis is
different. This plant is peculiar to Western Australia. In the course of some
years I have received humerous specimens—most of them from Lieut.-Col. B. T.
Goadby. I have been unable to detect the slightest difference in the floral
structure from that of P. nana. But the type form of P. pyramidalis has no
basal rosette, and well-formed stem leaves are present alternating up the stem.
This gives it a very distinctive appearance, and compels me to place it far from
P. nana, in spite of the complete identity of floral structure. Dr. Rogers considers
it a larger plant than P. nana, but there are records of dimensions of the latter
not far behind those he gives for P. pyramidalis. P. nana is also in Western
Australia, and I have received intermediate forms with both basal and cauline
leaves. It seemed to me therefore that P. pyramidalis should be treated as a
Western Australian variant of P. nana, its foliation constituting an exception to
the rule of Section A, Subsection a. This view has the support of some of my
fellow-workers in orchid study, but Dr. Rogers, who has seen and collected
P. pyramidalis in its native haunts, considers it wiser to keep these two separate.
With regard to P. Suttonii—a small, solitary specimen reached me some years
ago from Tasmania with this name attached. The Ven. Archdeacon Atkinson,
who sent it, was uncertain of the founder of the species, and all efforts to discover
any reference to the name have failed. I do not feel justified, therefore, in
including it.

BY REV. H. M. R. RUPP. 423

The following two notes affect certain groups in the classification:

(1) Section A, Subsection «a.—Exception to description of Subsection: P.
alpina Rogers.—No radical rosette or cluster of leaves has been observed in this
species: but it is so obviously closely allied to its neighbours that I prefer to
leave it with them and note the exception. Mr. EH. HE. Pescott (Orchids of
Victoria, p. 82) would apparently include P. furcata Lindl. in the same category,
but my own experience of the species does not confirm this (see also Plate VIII
in Rogers, Notes on Certain Species of Pterostylis, Proc. Roy. Soc. Victoria,
Nov., 1915).

(2) Section A, Subsection a, Division I. ‘Stem leaves reduced to bracts.’’—
When these plants occur in long grass or undergrowth, there is a tendency for
the radical rosette to disintegrate, the leaves ascending the stem towards the
light. This is very noticeable in P. nutans R. Br. and P. Baptistii Fitzg.; and
I am confident that Fitzgerald’s type specimens of the latter must have grown
under such conditions, for the foliation depicted by him is quite unusual. (When
it was described, the species was known only in one locality.) The normal
habit of P. Baptistii is undoubtedly to produce a very definite and fairly compact
radical rosette, with merely 2-3 more or less appressed bracts on the stem. In
the young stage its rosette is very similar to that of P. pedunculata R. Br.

It will be observed that I include P. Daintreyana F.v.M. in a Catochilus
group, not in Antennaea as Bentham does. This treatment is already, I think,
endorsed by general consent of all who know the species.

PTEROSTYLIS R. Br.

Section A. Laminatae.—Labellum laminate, from broadly oblong to linear, usually with
entire margins but occasionally cleft or emarginate at the apex, the lamina and
margins sometimes beset with translucent setae. Galea curving forward.

Section B. Filiformae.—Labellum filiform-terete, beset with long yellow hairs nearly
to the apex, where it terminates in a dark, knob-like appendage. Galea erect.

Section A.

Subsection A.—Radical leaves in a rosette or cluster encircling the base of the stem.
Stem leaves reduced to sheathing bracts or scattered along the stem. [Exception:
P. alpina. See above, Note (1).]

Division I.—Stem leaves reduced to bracts. [See above, Note (2).]
i. Flower solitary (rarely 2), sometimes large. Lower sepals erect.

iG 1. Labellum bifid. Bracts 2, or reduced to the one under the ovary.—
= P. ophioglossa R. Br., P. concinna R. Br.

4 2. Labellum entire. Bracts 2 or 3.

2 a. Labellum linear-lanceolate, decurved.—P. acuminata R. Br., P. Baptistii
S Fitzg., P. curta R. Br., P. nutans R. Br., P. Vereenae Rogers, P. depauperata
= Bailey.

& b. Labellum oblong, nearly straight.—P. nana R. Br. [but see note above on
E P. pyramidalis], P. pedoglossa Fitzg., P. pedunculata R. Br.

ii. Flowers racemose or spicate, seldom large. Lower sepals deflexed or spreading.
ef 1. Labellum shortly oblong, entire, bare. Sepals short and blunt.—P. mutica
5 R. Br., P. cycnocephala Fitzg.

Biel 2. Labellum ovate-oblong, sometimes emarginate, with a few long marginal
38 setae and very small crowded laminal ones. Sepals acuminate or prolonged
& into filiform points.—P. rufa R. Br., P. pusilla Rogers, P. Mitchellii Lindl.,

P. squamata R. Br., P. Woollsii Fitzg.

Division II.—Stem leaves well developed, scattered. Flower solitary (rarely 2),
often large. Lower sepals erect.

i. Labellum minutely bifid. Radical leaves 2-4.—P. furcillata Rupp.

ii. Labellum entire. Radical leaves many or few, rarely absent.—P. cucullata R. Br.,

P. falcata Rogers, P. alpina Rogers [see note above], P. furcata Lindl.,
P. gracilis Nicholls.

An-
tennaea

424 THE GENUS PTEROSTYLIS,

Subsection B.—Radical leaves in a rosette usually independent of the stem, occasionally
attached to it by a scape, but rarely encircling it, and often absent at flowering
time. Stem leaves at the base reduced to minute bracts, above usually well developed
and regularly alternate: occasionally bract-like.

Division I.—Flower solitary (rarely 2), sometimes large. Lower sepals erect with
long filiform points.
i. Labellum clavate at the apex.—P. grandiflora R. Br., P. constricta Sargent.
ii. Labellum minutely bifid.—P. Toveyana Ewart and Sharman.

8 iii. Labellum entire [occasionally bifid in P. Rogersii].

Ss 1. Labellum acute or acuminate [occasionally obtuse in P. truncata].—P. alata

a Reichb. f., P. robusta Rogers, P. Rogersii Coleman, P. revoluta R. Br.,

r= P. refleca R. Br., P. coccinea Fitzg., P. truncata Fitzg.

< 2. Labellum obtuse.—P. obtusa R. Br., P. decurva Rogers, P. recuwrva Benth.,
P. pyramidalis Lindl.

Division IIl.—Flowers racemose (rarely solitary), very small. Lower sepals erect,
very short. Rosette sometimes encircling the stem.—P. parviflora R. Br.
(Antennaea. )

Division III.—Flowers racemose (rarely solitary), seldom large. Lower sepals
deflexed or spreading.

on i. Labellum deeply trifid.—P. Sargentii Andrews.
me ii. Labellum entire or emarginate.—P. Daintreyana F.v.M., P. longifolia R. Br.,
og P. vittata Lindl.

Section B.—Flower usually solitary, often rather large. Lower sepals more or iess
deflexed. Leaves radical and cauline, only slightly spreading.—P. barbata Lindl.,

P. turfosa Endl. (Catochilus.)

The distribution of these 46 species is interesting. Only one*—P. rufa R. Br.—
is recorded from every State, and in view of the considerable confusion between
this species and its nearest allies, I think it is more than likely that some of the
records should be transferred to P. pusilla, P. Mitchellii, or P. squamata. The
distinctions are admirably set out on p. 42 of Dr. Rogers’ “South Australian
Orchids”. There are, however, certain forms frequently included in P. pusilla
which do not quite conform to Dr. Rogers’ descriptions of that species, being
more robust, carrying more flowers, and having simpler details in the labellum.
One of these—a New South Wales form—I have named var. prominens (Proc.
Linn. Soc. N.S.W., lvi, 1931). This is linked up with the type by certain inter-
mediate forms in southern States. In the table of distribution which follows, I
have queried the records of P. squamata in Tasmania, because I have reason to

* Since this paper was read I have been informed that P. nutans has been
discovered in Western Australia, and if this be so it also occurs in all States.

A. Examples of labellum in Section A (Laminatae).

B. Labellum in Section B (Filiformae).

1-4. Foliation and inflorescence in Section A, sub-section A. 1-2, Division I, i (P. nutans,
P. ophioglossa) ; 3, Division I, ii (P. pusilla v. prominens) ; 4, Division II (P. falcata).

®-7. Foliation and inflorescence in Section A, sub-section Bs. 5-6, Division I (P: reflexa,
P. truncata); 7, Division III (P. longifolia).
(Division II is omitted, as it contains but one species, with the smallest flowers in
the genus.)

8. Foliation and inflorescence in Section B (P. twrfosa).

9-31. Labella, front view (flattened out), of various species in Section A.—9, P. ophio-
glossa; 10, P. concinna; 11, P. acuminata; 12, P. Baptistii; 13, P. curta; 14,
P. pedunculata; 15, P. nana; 16, P. mutica; 17, P. cycnocephala; 18, P. pusilla;
19, P. rufa; 20, P. Witchelli; 21, P. furcillata; 22, P. revoluta; 28, P. cucullata;
24, P. grandiflora; 25, P. Toveyana; 26, P. alata; 27, 27a, P. Rogersii; 28, P. obtusa;
29, P. parviflora; 30, P. Daintreyana; 31, P. longifolia.

32, 338, erect lower sepals.

34, 35, 36, deflexed lower sepals.

BY REV. H. M. R. RUPP. 425

If ( {3 |4 j IG {7

D7 29 23 24 25 26 Lp 2a 723.29 30 3

ae

Text-figures A, B, 1-36.
Genus Pterostylis R. Br.—Figures of Australian Species illustrating the classification

e
proposed in this paper.

426 THE GENUS PTEROSTYLIS,

believe that P. pusilla should replace it. I have collected the last-named near
Launceston: the specimens agree precisely with a Tasmanian specimen in the
National Herbarium at Melbourne, endorsed by F. v. Mueller as P. squamata;
but if Fitzgerald interprets the latter rightly it is impossible to accept this deter-
mination, and my specimens are almost typical P. pusilla. Two other records I
have marked as doubtful—P. cucullata in New South Wales and P. reflexa in
Queensland. While I think it quite likely that the former may occur in southern
New South Wales, all the specimens under that name which I have inspected
must be referred to a very distinct species, Dr. Rogers’ P. falcata. Bailey’s
description of P. reflexa (Q’land Flora) seems to me to indicate P. revoluta, which
he regards as conspecific; but a wide experience of both forms in New South
Wales has convinced me that Brown was quite right in separating them. I have
omitted Tasmania for P. obtusa, as all the Tasmanian plants under this name,
which I have collected or seen, are typical specimens of the allied species
P. decurva. About P. depauperata little seems to be known, but I have retained it
because Bailey’s description appears to indicate a valid species, and it is quite
likely that a plant apparently not recorded south of Cairns should be distinct
from southern forms.

Seven species occur in all States except Western Australia, viz—P. concinna,
P. curta, P. nutans, P. pedunculata, P. mutica, P. parviflora, P. longifolia. Two
occur in all States except Queensland: P. nana and P. barbata. Four are restricted
to New South Wales, Victoria, and Tasmania: P. pedoglossa, P. falcata, P. alpina,
P. decurva. Except Victoria and Tasmania, each State has one or more species
not known at present in any other State—Queensland: P. depauperata; New
South Wales: P. clavigera, P. furcillata, P. coccinea, P. Daintreyana; South
Australia: P. Vereenae; Western Australia: P. Rogersii, P. constricta, P. recurva,
P. Sargentii, P. turfosa. P. pyramidalis.

The total number of species for each State (excluding those marked doubtful)
is as follows: New South Wales 33, Victoria 32, Tasmania 22, South Australia 21,
Queensland 16, Western Australia 11.

Abbreviations used in the Table of Distribution are:

Fitzg. = Fitzgerald. Australian Orchids, Vol. i or ii, and number of part.

Benth. = Bentham, Flora Australiensis, Vol. vi.

Q. Fl. = Bailey, Queensland Flora.

Intr. S.A. Orch. = Rogers, Introduction to the Study of South Australian
Orchids.

S.A. Orchids = Rogers, South Australian Orchids.

Orch. Vic. = Pescott, Orchids of Victoria.

Orch. N.S.W. = Rupp, Guide to the Orchids of New South Wales.

W.A. Orch. = Mrs. E. Pelloe, Western Australian Orchids.

Vict. Nat. = The Victorian Naturalist.

Linn. Soc. N.S.W. = PRocEEDINGS oF THE LINNEAN Socrtety or New SoutH
WALES.

Royal Soc. Vict. = Proceedings of the Royal Society of Victoria.

Royal Soc. 8. Aust. = Proceedings of the Royal Society of South Australia.

Royal Soc. W. Aust. = Proceedings of the Royal Society of Western Australia.

Journ. W.A. Nat. Hist. Soc. = Journal of the Western Australian Natural
History Society.

An asterisk (*) denotes that the reference is primarily to an illustration.

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POS US OU aU IT

BY REV. H. M. R. RUPP. 427

DISTRIBUTION OF AUSTRALIAN SPECIES.

With References to Descriptions or Illustrations,

. ophioglossa R. Br. Q’land, N.S.W. (Fitzg., I, 6*; Orch. N.S.W., 99*).

var. collina Rupp. N.S.W. (Linn. Soc. N.S.W., liv, 1929*).
concinna R. Br. Q’land, N.S.W., Vict., Tas., S. Aust. (Fitzg. I,-6* ; Orch. N.S.W., 100*).
acuminata R. Br. Q’land, N.S.W., Vict. (Fitzg. I, 5*; Vict. Nat., Aug., 1926, 102*).
var. ingens Rupp. Vict. (Linn. Soc. N.S.W., liii, 1928).
Baptistii Fitzg. Q’land, N.S.W. (Fitzg., I, 1* (see note above); Orch. N.S.W., 102%).
curta R. Br. Q’land, N-:S:iw., Vict., Tas., S. Aust. (Fitzge., I, 5*; Intr. S.A. Orch., 17*).
nutans R. Br. Q/land, N.S.W., Vict., Tas., S. Aust. (Fitze., I, 6*; Orch. N.S.W., 103%).
var. hispidula R. Br. Q’land, N.S.W. [Others?] (Fitzg., I, 6*).
Vereenae Rogers. S. Aust. (S.A. Orchids, 39).
depauperata Bailey. Q’land (Q. Fl., 1577).
clavigera Fitzg. N.S.W. (Moore and Betche, Flora of N.S.W., p. 400).
NANG Re Bre ONeSswe, Vict., as: Si Aust, W. Aust. (intr: S’A. Orch., 13*; Vict: Nat.,
Aug., 1926, 106*).
pedoglossa Fitzge. N.S.W., Vict., Tas. (Fitzg., I, 3*; Vict. Nat., July, 1926, 74*).
pedunculata R.Br. Q’land, N.S.W., Vict., Tas., S. Aust. (Fitzg., I, 5*; Vict. Nat.,
July, 1926, 74*).
mutica R. Br. Q’land, N.S.W., Vict., Tas., S. Aust. (Fitzg., I, 2*; Intr. S.A. Orch., 16*).
cycnocephala Fitzg. N.S.W., Vict., Tas., S. Aust. (Fitzg., I, 2*; Vict. Nat., July,
1926, %1*).
rufa R. Br. Q’land, N.S.W., Vict., Tas., S. Aust., W. Austs« (Fitzg., I, 2*; Vict. Nat.,
Aug., 1926, 106*).
pusilla Rogers. N.S.W., Vict., Tas., S. Aust. (Royal Soc. S. Aust., xlii, 1918*; Vict.
Nat., Aug., 1926, 106*).
var. prominens Rupp. N.S.W. (Linn. Soc. N.S.W., lvi, 1931).
Mitchellii Lindl. Q’land, N.S.W., Vict., S. Aust. (Vict. Nat., Aug., 1926, 106*; S.A.
Orchids, 42).
Squamata R. Br. N.S.W., Vict., Tas. ?, S. Aust. (Fitzg., I, 6*; S.A. Orchids, 42).
Woollsiit Fitzg. N.S.W., Vict. (Fitzg., I, 2*; Vict. Nat., Dec., 1928, 223*).

. furcillata Rupp. N.S.W. (Linn. Soc. N.S.W., lv, 1930, p. 415).
. cucullata R. Br. N.S.W. ?, Vict., Tas., S. Aust. (Royal Soc. Vict., xxviii, n. ser.,

iParntel*) Orch. Vici, i32*))

. faleata Rogers. N.S.W., Vict., Tas. (Royal Soc. Vict., l.c.*; Vict. Nat., July, 1926,

Ta) ye

alpina Rogers. N.S.W., Vict., Tas. (Royal Soc. Vict., l.c.*; Viet. Nat., l.c.*).
furcata Lindl. N.S.W., Vict., Tas., S. Aust. (Royal Soc. Vict., l.c.*; Vict. Nat., Aug.,
1926, 106*).

gracilis Nicholls. Vict., Tas. (Vict. Nat., March, 1927, 324*).

grandiflora R. Br. Q’land, N.S.W., Vict. (Vict. Nat., Aug., 1928, 112*).

constricta Sargent. W. Aust. (Royal Soe. W. Aust., 4, 1907; W.A. Orch.).
Toveyana Ewart & Shar. Vict., Tas. (Vict. Nat., July, 1926, 74*).

alata Reichb. f. N.S.W., Vict., Tas., S. Aust. (Fitzg., I, 3*, as P. striata; Vict. Nat.,
Aug., 1926, 102*).

robusta Rogers. Vict., S. Aust., W. Aust (Royal Soc. S. Aust., li, 1927; Intr. S.A. Orch.,
8*, as P. reflexa).

Rogersii Coleman. W. Aust. (Vict. Nat., Sept., 1929, 100*; W.A. Orch.*).

revoluta R. Br. Q’land, N.S.W., Vict. (Fitzg., I, 5*, in P. reflexa; Orch. Vic., foll.
Dp 32).

refleca R. Br. Q’land ?, N.S.W. (Fitzg., I, 5*; Orch. N.S.W., 110*).

coccinea Fitzg. N.S.W. (Fitzg., I, 6*; Orch. N.S.W., 112*).

truncata Fitzg. N.S.W., Vict. (Fitzge., I, 4*; Orch. N.S.W., 109*).

obtusa R. Br. Q’land, N.S.W., Vict., S. Aust. (Fitzg., I, 6*; Orch. N.S.W., 113*; Vict.
Nat., July, 1926, 74*).

decurva Rogers. N.S.W., Vict., Tas. (Royal Soc. S. Aust., xlvii, 1923*; Orch. Vic.,
foll. p. 48*).

recurva Benth. W. Aust. (Benth., 360; Fitzg., II, 2*; W.A. Orch., 64*).

pyramidalis Lindl. W. Aust. (Benth., 357; W.A. Orch.).

parviflora R. Br. Q’land, N.S.W., Vict., Tas., S. Aust. (Fitzg., I, 7*; Vict. Nat., July,
1926, 74*).

Sargentii Andrews. W. Aust. (Journ. W.A. Nat. Hist. Soc., May, 1905; W.A. Orch.*).

428 THE GENUS PTEROSTYLIS.

P, Daintreyana F.v.M. N.S.W. (Fitzg., I, 6*).
P. longifolia R. Br. Qjland, N.S.W., Vict., Tas., S. Aust. (Fitzg., I, 1*; Intr. S.A.
Orch T4)e
P. vittata Lindl. Vict., Tas., S. Aust., W. Aust. (Intr. S.A. Orch., 13*; Vict. Nat., Aug.,
1926, 102*).
var. subdifformis Nicholls. W. Aust. (Vict. Nat., Feb., 1933, 253).
var. viridiflora Nicholls. W. Aust. (Vict. Nat., l.c.).
P. barbata Lindl. N.S.W., Vict., Tas., S. Aust., W. Aust. (Fitzg., I, 7*; Orch. Vic.,
foll. p. 32*).
P. turfosa Endl. W. Aust. (Fitzg., II, 2*; W.A. Orch., p. 64*).

I am much indebted for valuable notes and suggestions in connection with
this paper to Dr. R. S. Rogers, Adelaide; Mr. EH. Nubling, Sydney; Mrs. C. A.
Messmer, Lindfield; Mr. W. H. Nicholls, W. Footscray (Vic.); and Mrs. Hdith
Coleman, Blackburn (Vic.). The articles in the Victorian Naturalist for July
and August, 1926, referred to so often above, are by Mr. Nicholls, who figures
there all the Victorian species.

A NEW SPECIES OF PTEROSTYLIS.
By (Mrs.) PrEArt R. MESSMER.

(Ten Text-figures. )

[Read 29th November, 1933.

PTEROSTYLIS PULCHELLA, Nl. Sp.

Planta gracilis 7-17 cm. alta, folius radicalibus rosulatis ad caulis basem scapo
affixis, saepe absentibus: folia superiora alternia, lanceolata. Flos solitarius circa
galea flexum 4-43 cm. longus, infra translucens lineis viridibus, supra rufus.
Galea breviter filamentosa, non deflexa. Petala paena tam longa quam sepalum
dorsale infra lata, supra concava; angustiana usque ad puncta maxime acuminata.
Sepala inferiora infra lata, aliquanto gibbosa ad sinum, supra subito filamentosa,
galeam multo excedentia. Labellum linearis-oblongum 14-17 mm. longum ad
apicem mutabiliter bifidum rufum atrum. Appendix fasciata 5 mm x 1 mm. ad
apicem dentata, dens media longissima. Columna labello fere brevior cum alis
glabris; anguli superiores acutissimi vex superiores quam antheri, stigma longum,
angustum, obscurum.

A slender plant 7-17 cm. high. Radical leaves in a rosette attached by a
scape to the base of the stem, often absent at time of flowering. Stem leaves
alternate, narrow to broad-lanceolate, sheathing at base. Flower solitary, 4-43 cm.
long round the curve of the galea, translucent-white with green bands below,
red-brown above. Galea curving gradually up to the point where it is embraced
by the lower sepals, thence horizontal to the finely acuminate or shortly filamentose
apex. Petals nearly as long as the dorsal sepal, broad below, concave above and
narrowing to finely acuminate points. Lower sepals broad below, somewhat
gibbous at the sinus, above suddenly contracted into filaments extending far
above the galea. lLabellum linear-oblong, 14-17 mm. long, variably bifid at the
apex, dark red. Appendage fasciate, 5 mm. x 1 mm., deeply dentate at the apex,
the median tooth the longest. Column a little shorter than the labellum, wings
glabrous with very acute upper angles hardly exceeding the height of the anther.
Stigma long, narrow, obscure.

The first specimens of this plant were discovered in April, 1932, in a parcel
of material sent to me for examination by Mr. R. MecNall, who found them
growing on rocks and associated with Liparis reflexa, Hymenophyllum
Tunbridgense and Dicranoloma Billardieri in the moist shade of the gorge of
Fitzroy Falls, N.S.W.

Though like no other known Pterostylis, it suggests affinities with P. grandi-
jfiora, P. ophioglossa and P. reflera, and may even have originated as a natural
hybrid between P. grandiflora and P. ophioglossa as the first specimens examined
suggested; but more material being available this year (1933) has placed it in
the rank of a constant species. The distinguishing features, though variable, are

I

430 A NEW SPECIES OF PTEROSTYLIS.

constant in their distinction, i.e., the shape and set of the apex of the galea, the
labellum and the appendage at the base of the labellum. In dissection the dorsal
sepal approaches that of P. ophioglossa, the lower sepals those of P. grandiflora.
The column appears to be intermediate between that of P. ophioglossa and that
of P. grandiflora, whilst the petals and base of the labellum are nearer to those

of P. reflexa.

Text-figs. 1-10.

Pterostylis pulchella, n. sp.
1.—Plant with obtuse leaves and flower having an obtuse subtending bract.
2.—Plant with very acute leaves. 3, 4.—Variations in leaf. 5.—Dorsal sepal.
6, 7.—Petals acuminate and shortly filamentose. 8.—Lateral sepals. 8a.—Showing
variations in sinus between lateral sepals. 9, 9a.—Labella. 10, 10a.—Columns.
Fig. 1 is of the type specimen.

THE MYCETOZOA OF NEW SOUTH WALES.
By Litisn FrAseER, M.Sce., Linnean Macleay Fellow of the Society in Botany.

[Read 29th November, 1933.

The members of the class Mycetozoa or Myxomycetes have a remarkably
wide distribution throughout the world. They are found both in tropical and in
cold regions, but appear to be most abundant in the cooler moist parts of the
temperate zone.

Many species are known which are common in Hurope, America, Australia,
ete., and which retain their specific characters exactly in all localities. On the
other hand, certain species vary widely in the same locality, and even in the
one group of individuals.

The wide distribution of the class as a whole, and of individual species in
particular, is probably due to the fact that their spores are very small and light,
and are therefore readily transportable. Also the spores of some species are
known to remain in a viable condition for as long as 7 years. Most, however,
will not germinate after 1—2 years.

The members of the class Mycetozoa are chiefly small and insignificant, so
that they must frequently be overlooked by collectors. Also their habitat is, as
a rule, inconspicuous. Principally, they are found on decaying fallen logs and
bark, frequently on the lower surfaces, and on dead leaves and grass. Some are
cosmopolitan in their habitat, for example the Physarums, which are found on
wood, grass, and leaves. Others are more restricted in their distribution.
Cribraria and Dictydium, for example, occur on decaying wood. Fuligo septica,
on the other hand, is widely distributed; it is common on the sawdust in tan-
yards, on decaying wood and bark, on twigs, leaves, and grass. Some species of
Comatricha and Stemonitis are found only on wood or bark, others are chiefly
found on leaves.

Cooke (1892) records 49 species of Mycetozoa collected from Australia, but
of these only Arcyria Oerstedtii Rost. (A. fuliginea Mass.), Mucilago spongiosa
Morgan (Spumaria alba Bull.), and Badhamia varia Mass., which has been
subdivided into three species by Lister (1925), are reported specifically from
New South Wales. Three additional species are given by McAlpine (1895), but
of these none is from New South Wales. A later list by McAlpine (1898) includes
Physarum cinereum Pers., Stemonitis fusca Roth., and Badhania utricularis Berk.

In 1918, Cheel published a list comprising those species already known from
New South Wales, together with a large number of new records. This list
contained 48 species, representing 28 genera and 11 families. The next important
contribution was by Cleland (1927), the number of species recorded from New
South Wales being raised by 12. Cleland’s record also furnishes data as to the
relative abundance of the species occurring here.

In the following list, 23 species are recorded for the first time from New
South Wales. The species already reported by the authors mentioned, are also

432 THE MYCETOZOA OF NEW SOUTH WALES,

incorporated in the list. Notes on the habitat, abundance, and seasonal occurrence
are given wherever possible.

1. Subclass EXosporEAr.
Family CERATIOMYXEAE.

Ceratiomyza fruticulosa Macbr. One of the commonest of the Mycetozoa,
found on the coast and highlands throughout the year, especially during the
autumn, on rotting wood and bark. (C., Cl., F.)*

2. Subclass ENDOSPOREAE.
Family PHYSARACEAE.

Badhamia capsulifera Berk. Orange, October, 1916. (Cl.)

B. papaveracea Berk. & Rav. Sydney district, on apple twigs, March, 1931. (F.)

B. utricularis Berk. (McAIp., C.)

Physarum leucopus Link. Sydney district, on earth, leaves and sticks, May,
UBL | (aE))

P. dictyospermum Lister. (C.)

P. viride Pers. One of the commonest of the Mycetozoa on the coast and
highlands. Found ai all times of the year, on dead wood, bark, twigs and
leaves. (C., Cl., F.)

P. viride var. aurantium Lister. Sydney district, on dead wood, March, 1931.
Mt. Wilson, on dead wood, March, 1931. (F.)

P, viride var. incanum Lister. Fairly common in the Sydney district, especially
during the autumn, on dead wood. (C., F.)

P. rigidum G. Lister. Mt. Wilson, on dead wood and bark, March, 1931. Sydney
district, on wood, March, 1931. (F.)

P. flavicolum Berk. Fairly common in the Sydney district, on dead wood.
(Lister, C., F.)

P. Maydis Torrend. Sydney district, on dead log, April, 1931. (F.)

P. pusillum Lister. Sydney district, on dead bark and wood, March, 1931. (F.).

P. didermoides Rost. Wollongbar, N.S.W. (Cl.)

P. nutans Pers. Common around Sydney, on dead wood. (C., Cl. F.)

P. nutans var. leucophaeum Lister. Common in the Sydney District. (C.)

P. compressum Alb. & Schw. Sydney district, August. (Cl.)

P. reniforme Lister. Hawkesbury River, Wollongbar, June, 1913. | (Cl.)

P. cinereum Pers. Common throughout New South Wales, especially in the
autumn. (C., Cl., F., McAlp.)

P. vernum Somm. Mt. Wilson, on dead wood, March, 1931. Sydney, on

twigs, straw, etc., October, 1930. Warialda, on grass, December, 1931. (F.)

P. gyrosum Rost. Sydney Botanic Gardens. (C.)

P. sinuosum Weinm. Sydney district. (C.)

P. bitectum Lister. (C.)

Fuligo septica Gmelin. Common throughout the State, especially the coast
and highlands, at all times of the year, but mostly in the summer, on
wood, twigs, ete. (C., Cl. F.)

F. septica var. candida Fr. Broken Hill, April, 1917. Hawkesbury River,
January. (Cl.)

F. cinerea Morgan. Hawkesbury River, April, 1913. Sydney district, March,
1914. (Cl.)

=(€. = Cheel, Cl: = ‘Cleland, B= Eraser, McAlp. = McAlpine:

BY LILIAN FRASER. 433

Cienkowskia reticulata Rost. Sydney district, on grass, May, 1931. Mt. Wilson,
on dead wood and bark, March, 1931. (F.)

Craterium leucocephalum Ditm. Sydney district, on dead leaves and twigs,
March, 1931. (F.)

C. leucocephalum var. scyphoides Lister. Sydney district, on twigs, May, 1931.
(F.)

Leocarpus fragilis Rost. (C.)

Diderma effusum Morgan. (C.)

D. radiatum Lister. (C.)

Diachaea leucopoda Rost. Common around Sydney. (C., Cl.)

Family DipyMtraceaAr.

Didymium difforme Duby. (C.)

D. melanospermum Macbr. Sydney district, on dead twigs, March, 1931. (F.)

D. nigripes Fries. Sydney district and National Park, on dead leaves and
twigs, not uncommon. (Cl., F.) Specimens near to D. nigripes var.
xanthopus Lister are also not uncommon.

D. squamulosum Fries. (C.)

D. leoninum Berk. & Br. (C.)

Mucilago spongiosa Morgan. (Cooke, C.)

Family CoLLoDERMACEAE.
Colloderma oculatum G. Lister. (C.)

Family SreEMONITACEAE.

Stemonitis fusca Roth. Fairly common and widespread, especially in autumn,
on dead wood. (McAlp., C., F.)

S. splendens Rost. Common and widespread throughout the State at all times
of the year, on dead wood and bark. (Cl. F.)

S. splendens var. Webberi Lister. National Park. (C.)

S. splendens var. flacida Lister. Canberra, January, 1933. (N. White.)

S. herbatica Peck. Sydney district, March, 1914; October, 1916. (C.)

S. ferruginea Ehrenb. Sydney district, on dead wood, March, 1931; July,
1932> (Parramatta. (C., E)

Comatricha nigra Schroet. Common throughout the State, especially in the
Sydney district, at all times of the year, on dead wood and bark. (C.,
Cl., F.)

C. lara Rost. Sydney district, on dead wood and bark, March, 1931. (F.)

C. elegans Lister. Sydney district, on dead wood and bark, July, 1932. (F.)

C. pulchella Rost. var. fusca Lister. Sydney district, March, 1931. (F.)

C. typhoides Rost. Fairly common in the Sydney district, on dead wood.
(Ge, Clk, 135)

C. irregularis Rex. (C.) :

Enerthenema papillatum Rost. Sydney district, on charred wood, March, 1931.
(F-)

Clastoderma Debaryanum Blytt. I have only collected this species once, at
Avoca, Gosford District, September, 1930. Rare. (C., F.)

Family HrrrropERMACEAE.
Cribraria argillacea Pers. (C.)
C. vulgaris Sehrad. Sydney district, on dead wood, April, 1932; National
Park, on dead wood, January, 1931. (F.)

434

THE MYCETOZOA OF NEW SOUTH WALES,

C. macrocarpa Schrad. Mt. Wilson, on dead wood, November, 1931. (F.)

C. intricata Schrad. var. dictydioides Lister. Sydney district, on dead wood,
March, 1931. (F.)

OC. tenella Schrad. var. concinna G. Lister. Sydney district, on dead wood,
May, 1931. Hawkesbury River district, on dead wood, July, 1932. (F.)

Dyctidium cancellatum Macbr. Sydney district, on dead wood, March, 1931.
(C22E4)

Family TUBULINACEAE.

Tubifera ferruginosa Gmel. Common throughout the coast and highlands, on
dead wood and bark. (C., Cl., F.)

Family R&ricULARIACEAE.
Dictydiaethalium plumbeum Rost. Common throughout the State at all times
of the year, on bark. (C., F.)
Reticularia lycoperdon Bull. Sydney district, May, 1918. (Cl.) Mt. Wilson,
on dead wood, March, 1931. (F.)

Family LycoGALAcEAE.
Lycogala epidendrum Fries. Common on the coast and highlands, especially
in the autumn, on dead wood. (C., Cl., F.)

Family TRICHIACEAE.

Trichia verrucosa Berk. Common on the coast and highlands, throughout
the year, on twigs, bark, wood, ete. (C., Cl., F.)

T. affinis de Bary. Fairly common in the Sydney district, on dead twigs,
leaves and wood. (C., F.)

T. persimilis Karst. (C.)

T. varia Pers. Very common on the coast and highlands, especially about
Sydney, throughout the year, on dead twigs, leaves and wood. (C., Cl., F.)

TNenecta sRexan (C3)

T. decipiens Macbr. Common throughout the year, on dead wood. (C., F.)

T. floriformis G. Lister. Very common throughout the year in the Sydney
district, and highlands, on dead wood. (C., Cl., F.)

Henvitrichia Vesparium Macbr. Sydney district on dead wood, February, 1931.
(Gigg: 185))

H. clavata Rost. Sydney district, on charred wood, July, 1932. Mt. Wilson,
on dead wood, March, 1931. (F.)

H. Serpula Rost. (C.)

Family ARCYRIACEAE.

Arcyria ferruginea Sauter. Coastal district and mountains, very common
throughout the year. (C., Cl., F.)

A. cinerea Pers. Common in the Sydney district, throughout the year, on
dead wood. (C., Cl., F.)

A. denudata Sheldon. Common in the coastal district and mountains through-
out the year, on dead wood. (Cl., F.)

A. insignis Kalchbr. & Cooke. (Cl.)

A. glauca Lister. Sydney district, on dead twig, May, 1931. (F.)

A. incarnata Pers. Botanic Gardens, Sydney. (C.)

A. incarnata var. fulgens Lister. Mt. Wilson, on dead wood, March, 1931.
(F.)

BY LILIAN FRASER. 435

A. nutans Grev. Fairly common in the Sydney district, on dead wood.
(C., Cl.)

A. Oerstedtii Rost. On leaves of Atherospermum (Cooke).

Perichaena depressa Libert. Hawkesbury River. (Cl.)

P. corticalis Rost. (C.)

P. vermicularis Rost. Sydney district, on dead wood and bark, March, 1931.
(F.)

Family MARGARITACEAE.

Dianema depressum Lister. Mt. Wilson, on dead wood, August, 1932. (F.)

This list of Mycetozoa includes 88 species and varieties, most of them being
recorded from the environs of Sydney and the adjacent highlands. There is no
reason to doubt that they occur in equal numbers along the whole of the coast
and highlands, since the climate is quite suitable for their development. Very
few records are known for the western parts of the State, and it seems quite
probable that they do not occur there at all frequently, the relatively hot and
dry climate being unsuitable for their development.

In the coastal and mountain districts, the Mycetozoa are chiefly found along
the margins of shady creeks and similar moist places in and bordering patches
of rain forest. Few are reported from the open sunny woodlands, even of the
coastal district. Of those which have been found in relatively exposed places the
following are the most common: Arcyria ferruginea, Fuligo septica, Didymium
spp., Dictydiaethalium plumbeum. Dianema depressa and the various species of
Cribraria and Dictydium are found only in the most shady, moist places.

The genera in which most species have been recorded are: Physarum (19
species), Arcyria (9 sp.), Trichia (7 sp.), Comatricha (6 sp.), Stemonitis (5 sp.),
Cribraria (5 sp.), Didymium (5 sp.).

From the point of view of world distribution, as well as their numbers in
Australia, these genera are amongst the largest, in that the largest number of
different species have been attributed to them.

There are, however, several notable exceptions. The genera Badhamia,
Diachaea, Diderma, and Hemitrichia are poorly represented, there being only
3, 1, 2 and 3 species respectively reported from New South Wales, while no species
of the genus Lamproderma is known here. It may be that further search will
reveal more of these species, but at present it seems as if many common northern
hemisphere species are lacking here. There is also a significant lack of the
limey stalked Badhamias and Physarums. The warmer climate Physarums,
Cribrarias, Cienkowskia, etc., are, on the other hand, moderately well represented.

The commonest species amongst the collections so far made here are
Stemonitis splendens, Arcyria ferruginea, Ceratiomyxa fruticulosa, Physarum
viride and varieties, Dictydiaethalium plumbeum, Tubifera ferruginosa, Fuligo
septica, Trichia varia and T. floriformis. All of these are widespread, and most
of them are common throughout the world.

Only seven species of those recorded from New South Wales are not wide-
spread elsewhere. Of these, Physarum dictyospermum and P. flavicolum are
reported from New Zealand. Physarum Maydis, P. reniforme, P. gyrosum and
Didymium leoninum. are reported from Japan, Ceylon, Java, etc., and Arcyria
glauca has not been reported previously outside of Japan, where it has been
collected only on three occasions.

436 THE MYCETOZOA OF NEW SOUTH WALES.

In conclusion, I desire to thank Miss G. Lister, who identified some of the
species here recorded, and Professor T. G. B. Osborn, Department of Botany,
University of Sydney, for helpful advice.

Literature Cited.
CHEEL, E., 1918.—Notes on Australian Myxomycetes. Australian Naturalist, iv (1),

pp. 6-12.

CLELAND, J. B., 1927.—Notes on a collection of Australian Myxomycetes, identified by

Miss Gulielma Lister. Trans. Proc. Roy. Soc. Sowth Auwst., li, pp. 62-64.

Cooxs, M. C., 1892.—Handbook of Australian Fungi. London.
Lister, A., 1925.—A monograph of the Mycetozoa. Third Edition. Revised by G. Lister.

British Museum Publication.

McALPINE, D., 1898.—New Myxomycetes from New South Wales. Proc. LINN. Soc.

N.S.W., xxiii, pp. 82-84.

Postscript, added 30th November, 1933.—Dictydium rutilum G. Lister (Journ.
Bot., 1933, pp. 222-223) has recently been described from material collected in the
Sydney district, N.S.W., on decaying wood, February, 1931; and from Mt. Wilson,
on wood and bark, March, 1931. (F.)

THE COASTAL TABLELANDS AND STREAMS OF NEW SOUTH WALES.
By Frank A. Crart, B.Se., Linnean Macleay Fellow of the Society in Geography.
(Six Text-figures. )

[Read 29th November, 1933.]

Historical Introduction.

The use of distinctively physiographic methods in New South Wales is
virtually confined to this century, but modern conclusions were foreshadowed and
influenced by earlier work: Berry (1822) outlined the structural features of the
Sydney region and many surface relationships; Clarke (1853, 1860, 1875, 1876)
devoted much attention to stream origins, valley formation and denudation
generally; Thompson (1869) described erosional features in the Goulburn district,
and Wilkinson (1882) summarized the topographic development of the highlands.
The natural approach for observers was from the coast, so Berry (1825), Dana,
Jukes, Strzelecki and Wilkes (between 1845 and 1850) were impressed by
evidences of coastal movement, and sought to explain the precipitous faces of
plateaus fronting the ocean by faulting, and submergence of downthrown areas.
In 1845 Murchison referred to the eastern highlands of Australia as the ‘“‘Cordillera”
on the ground of geological resemblances to the Urals, and Wilkinson (1882)
continued the term, describing the cordillera as being originally a range of
upheaval whose sides had been shaped into deep valleys. He considered that its
highest points had probably not been completely submerged since the beginning
of the Mesozoic era, and postulated a Tertiary subsidence of the order of 4,000
feet over the eastern part of the continent to allow the development of the various
sediments of that age now found in the highlands. So far as external relationships
were concerned, Clarke held the Australian Cordillera and the New Zealand arc
to be similar anticlines separated by a trough, and to be comparable with the
ridges and inter-ridge valleys of such alluvial goldfields as Lachlan River.

Towards the end of the century, David established the basis for modern
discussion by giving further evidence of coastal movement (with Etheridge, 1890;
with Etheridge and Grimshaw, 1896b) and by proving the nature of the eastern
slope of the Blue Mountains (1896a, 1902). In the present century the study
of physiography as such was introduced by E. C. Andrews, who applied the
conclusions of North American workers to Australian problems and also con-
tributed to the theory of the subject; his principal writings dealt with the
Queensland coast and the eastern plateau (1902), New England (1903a, 1904a,
1905, 1912, 1914), the Blue Mountains (19030), Physical Geography of New South
Wales (19040), New Zealand Sounds (1906), floods (1907), the work of gravity
streams (1909), the geographical unity of Eastern Australia (1910a), Yosemite
(19100) and erosion (1911), while certain professional papers contained physio-
graphic references, especially those dealing with Hillgrove (1900), Kiandra (1901),

J

438 COASTAL TABLELANDS AND STREAMS OF NEW SOUTH WALES,

Drake (1908), Forbes-Parkes (1910c), and Cargo (1915). Other papers recognizing
similar principles were those of Jensen (1906, 1907), Sussmilch and Jensen (1909),
Sussmilch (1909, 1911, 1914, 1923), Woolnough and Taylor (1906), Taylor (1907,
1910, 1911, 1923a, 19230), and Hedley (1910, 1911). These all had certain aspects
in common; they adopted the idea of base-levelling, stream change by dynamic
action, the interpretation of surface history by the study of land forms, and a
time scale based on the physiographic position of Tertiary drifts and basalts,
and all had affinities with conclusions of the past century in respect to coastal
faulting, the cordillera, and the uniformity of movement in the eastern part of
the continent.

Earlier observers postulated extensive faultings because of the precipitous
coast and the sharp fall from a narrow continental shelf; later, Taylor held
similar views because he considered that streams rising near the coast and flowing
away from it (eg., Nepean tributaries) are abnormal, and must have been
balanced in the past by a further system to the east, where the sea is now;
he also required the suggested land mass as a premise for his views on the diversion
of former westerly and meridional drainage to the present east coast. This
gives a connection with the idea of the “cordillera’, for if that feature be con-
ceived as a meridional anticline, “the general east and west trend of the coastal
streams shows consequent drainage attendant on the seaward slope of the upland
during previous elevations’ (Andrews, 1902, p. 180), and meridional streams
require to be explained as ‘“subsequents” due to the expansion of original
“consequents” (Andrews, 1902, 19036, 1905), by meridional folding or faulting
attendant upon uplift (Hedley, 1911; Taylor, 1911, 1923), or by diversion through
block faulting in special cases (Sussmilch, 1909). At any rate, the idea of the
Cordillera involves a special explanation for the meridional streams, and has
helped towards the interpretation of many positive topographic features as being
of tectonic origin associated with the most recent (Kosciusko) warpings. This
view regards the Main Divide as being essentially an anticlinal crest, and
higher ridges parallel to it have been interpreted as former anticlinal divides,
as in the case of the heights of the Macleay basin (Andrews, 1903a, pp. 205-206),
although the effect of resistant rocks in the preservation of such high features
has also been noted (ibid., p. 196). This idea of moving anticlines was carried
to its greatest extreme by Taylor (19230), who supposed one of them to have
existed at a considerable height where the Sydney coast is now, and to have
subsided as the wave crest progressed westward .to form the existing Main
Divide (at 3,500 to 4,000 feet).

With respect to the question of unity or uniformity, it may be noticed that
Murchison (1845), Clarke (1876), and Wilkinson (1882) had the idea of uniformity
of structure and process in the whole of eastern Australia, and the last-named
envisaged a general rise or fall of much the same magnitude throughout the length
of the highlands (pp. 52-59). Andrews extended the idea (1910), especially on
the basis of widespread Tertiary peneplanation and subsequent differential uplift
in which the region had acted as a unit, although he recognized moderately high
relief in central New England following post-Cretaceous uplifts (1904, pp. 292-293):
similarly, Reid (1926) ascribed considerable relief to eastern Queensland
throughout the Tertiary period, and Craft (1932, 1933) regarded the plateau of
south-eastern New South Wales as having been relatively high over a similar
period. This emphasizes Andrews’ (1914) conclusion that Tertiary surfaces
were incipient peneplains not involving complete reduction, and puts the master

BY F. A. CRAFT. 439

peneplanation of the region back to late Palaeozoic or early Mesozoic times, with
decreasing age going northward, and some apparent diversity of features caused
by subsequent folding in Queensland as a variant of the gentle warping experienced
elsewhere.

However, the most distinctive features of modern thought are the recognition
of a peneplain now forming the principal surface of the plateaus, and the
theoretical conceptions associated with it. Thus Sussmilch (1911, p. 144) writes,
“The surface of the peneplain is not flat, but is traversed in most places by a
network of broad, shallow, mature valleys, ranging from 150 to 300 feet in depth”,
which are ascribed to the influence of a slight uplift (p. 146); David (1932, p. 172)
speaks of the same feature as having been “ one of the most perfect
peneplains imaginable; the slope of the land seawards being so slight as barely to
suffice for any run-off after rain’, and hills rising to similar heights above
the plain surface in various localities have been referred to as relics of older
regional peneplains (eg., Andrews, 1903a, pp. 216-217; Sussmilch, 1909,
pp. 335-344), without reference to their distance from the various divides. From
this it will be seen that the peneplain was conceived as a virtually horizontal
surface, and that successive peneplains had a generally similar form.

Now the present highlands offer considerably more variety of surface features
than this may indicate, but on account of the (supposed) general morphological
similarity of higher and lower surfaces, and their frequent separation by steep
slopes or rugged falling country, it has been inferred that a former common
surface has been broken up by block faulting and differential uplift, and that
the bounding slopes of the higher blocks are fault scarps (Sussmilch, 1909;
Andrews, 1910a). It must be remembered, however, that most high features are
near stream heads, or are composed of resistant rocks; that ‘“pre-fault’”’ surfaces
preserved by basalts show considerable local relief in many districts, ranging
from 700 to 1,400 feet in restricted areas without disturbance through more
recent uplifts, and that certain of the high blocks are bounded by normal
erosion escarpments (e.g., Shoalhaven, Craft, 1932), so an examination of plateau
topography begins without any preliminary assumption of extensive faultings.

Summing up, it has been found that certain ideas have been inherited from
early studies, namely, those of extensive coastal faulting, the origin of the
eastern highlands and the Main Divide in an anticlinal fold, and the uniformity
of process and movement in the history of the existing surface. Distinctively
modern conceptions are those of stream rearrangement by dynamic or tectonic
action, the formation of an approximately horizontal surface near sea-level, and
its elevation and breaking up to form the modern plateaus. The present work
bases no conclusions on the grounds of possible coastal faultings or movement;
it regards the Main Divide as a rather accidental feature of essentially non-
tectonic origin, and the river systems as having been stable during the present
physiographic record as the result of comparatively quiet evolution. A description
of surface features is attempted from a similar viewpoint.

TOPOGRAPHY.

Preliminary.—The plateau region consists essentially of three low domes—
New England, the Blue Mountains, and Monaro-Kosciusko—which are separated
from more northerly highlands and from one another by the Darling Downs,
Cassilis, and Lake George areas respectively. The middle part of each mass,
to a width of 50 miles from east to west, consists of a high plateau with streams

440

COASTAL TABLELANDS AND STREAMS OF NEW SOUTH WALES,

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Text-fig. 1.—Locality Map of the Region; the Main Divide is shown
by a heavy broken line. Names omitted from this map will be found on
Text-fig. 6.

BY F. A. CRAFT. 441

flowing considerably above sea-level: on either flank is a zone up to 50 miles in
width in which the streams fall abruptly towards sea-level, through gorges at
first, and then by way of widening valleys and plains. The major gaps of
Darling Downs and Cassilis slope away gently on either side, but there is no
break in the coastal heights east of Lake George, and streams flowing westward
(e.g., Lachlan River) from its vicinity pass through canyon sections. Beyond
the limits of the highlands are inland plains on the west, and the sea coast
with irregular, narrow plains on the east.

There is some variation of surface appearance in going from north to
south: thus the coastal slopes between the Queensland border and Hunter River
have rounded forms and many hills more or less isolated from the plateau
mass; the appearance of roundness is helped by forests and brush common to
the region. On the other hand, the coastal fall in the country south of the
Hunter consists mainly of solid ridges preserving even crests, with few isolated
hills. In the high plateau about Kosciusko all features in the region of heavy
rainfall are rounded, and on the western foothill slopes of Monaro-Kosciusko there
is a tendency towards isolated hills and concave slopes. However, a general rule
applies to the head parts of all the canyon sections, namely, outlines are
rectangular, with sharp breaks of slope, and with uniform slopes on the canyon
sides.

Exterior and Interior Forms.

In the higher plateaus and their slopes two broad types of scenery are
commonly presented. In the first, the whole countryside appears as an almost
horizontal plain, varied by occasional hills, and with any higher land at a distance,
generally on one or two sides only. In the second, there are plains of greater
or less extent terminated by superior walls or ridges, and generally appearing
aguite enclosed; the ridge summits give the impression of a continuous surface
when viewed from a point in or above their plane, but actually the whole ridge
features only occupy 10% to 25% of the locality, except where they expand into
plateaus, and their boundary is purely arbitrary. For the present discussion,
the terms “exterior” and “interior” will be applied respectively to the types: in
the former, the whole surface may be considéred as a unit without notable
break of slope, whether it is plain or undulating, while the latter depends on
the major break of slope in passing from central plains to bounding ridges, and
on the existence of well-differentiated planes inclined to one another. In addition,
the narrow canyons and the conical peaks of Barrington Tops at the head of
the Manning, Hunter and Peel Rivers are distinct scenic features. The principal
occurrences of the two main types may be summarized:

(a) Exterior Forns.
(i) Elevated Peneplains: New England; tableland of upper Macquarie-
Abercrombie Rivers; Kosciusko plateau; Monaro peneplain.
(ii) Surfaces of sub-horizontal Rocks: Plateaus of Hawkesbury and
eastern Shoalhaven; part of Clarence basin.
(6) Interior Forms.
(i) Ridge and Plain: Lower New England, Cassilis, and Liverpool Plains;
Goulburn-Lake George; Shoalhaven; Murrumbidgee; Federal
Territory; Western Monaro; Bathurst Plains.
(ii) Plateau and Plain: Hunter Valley; valleys of western Hawkesbury
(Kowmung, Cox, Colo, lower Wollondilly) ; coastal strip.

442 COASTAL TABLELANDS AND STREAMS OF NEW SOUTH WALES,

(a) Exterior Forms.

(i) Hlevated Peneplains.—The term “peneplain’” is used to denote those
surfaces in which erosion has been carried so far that original high watersheds
have been almost completely reduced, leaving occasional isolated hills or low
ridge lines. The land surface is composed of gently undulating country of low
relief, or of gently concave plains; the summit plane appears to be quite horizontal,
but on the falls from the central plateau mass it may be inclined at angles of
1° to 3°. The characteristic feature of the scenery is an even skyline, which
gives an appearance of horizontality that is often deceptive, as changing distance
and perspective mask undulations or gradual slopes over long distances. The
lower portions of the surface consist of valleys or plains, which are alluviated,
and support meandering normal streams that have flood terraces up to a mile
in width. Once the steep headwater portion of a stream is passed, a gentler
section is found at an almost constant range of altitude for each district, and
with a uniform depth below corresponding ridge crests. This is best shown on a
large scale in the Macleay basin, where radial streams and ridges fall at a
uniform rate towards a common centre over a distance of 20 miles. The continuity
of the upland features is interrupted by canyons of varying depth as the plateau
margins are approached, and the old peneplain surfaces are preserved in the
ridge crests and gentle upper slopes, until they become dismantled in the general
fall to lower country. There are some exceptions to this rule, as portions of
relatively undissected plateau come near the coast at intervals throughout its
length.

The principal areas comprised in the type are the upper surface of New
England above 3,000 feet, or above 2,000 feet to the north-west of Tenterfield;
the plateau of the southern Macquarie-Abercrombie-Jenolan region (mainly 3,000
to 4,000 feet); the country immediately west of Lake George (above 2,000 feet) ;
the Kosciusko-Kiandra-Upper Murray plateau (4,500 to 7,000 feet), and the
Monaro district (2,800 to 3,500 feet). In places the valleys in the edges of these
high blocks have been expanded into plains to form true interior features: chief
among these are the valleys of the Gwydir, Namoi and Peel Rivers (Liverpool
Plains), the Talbragar, Hunter, upper Murrumbidgee, and the Hucumbene-Snowy.
Such features, rather than the major differences in summit elevation between
the high-level peneplains, form the separation zones between the various plateau
groups.

(ii) Surfaces of sub-horizontal Rocks.—These are usually included with the
peneplains because slightly upturned edges have been bevelled off by erosion,
but they represent a distinct scenic type. As a general rule, they appear to be
within a few hundred feet of the original upper surfaces of deposition, but they
have been deformed into a series of shallow basins with inward slopes of the
order of 2°, which carry the surfaces from sea-level to a maximum altitude of
4,000 feet (near Lithgow). As with the elevated peneplains, there are no masses
enclosing the features, which are of a regional character, but there are sharp
rises from some edges to higher masses of the preceding class. Such edges are
found in the north and east of New England, facing the Darling Downs and the
Clarence basin respectively, and to the west of the Hunter-Sydney region, on a
line extending from Mudgee to Goulburn. The topography of these areas is
mainly controlled by elevation and rock character, because the bedding and
joints of the rocks give series of horizontal and vertical surfaces, thus imposing
a dominant convex profile in hill and valley sides, with minor cliffs at intervals,

BY F. A. ORAFT. 443

and a tendency towards major cliffs in the lower slopes, especially where softer
layers of rock are found. The skylines are very even, approximating closely
to the horizontal, with a few minor irregularities where basalt flows or ancient
monadnocks rise above the general surface. The major occurrences are on the
Darling Downs (Queensland), in the Clarence basin, the Hunter-Sydney region,
the upper Clyde, and the eastern Shoalhaven divides.

(b) Interior Forms.

In the type of exterior features, classification depends on the dominance of
one surface: with interior forms, a surface much below those high features of the
landscape is the criterion, and it is separated from them by a very marked
break of slope. There is no firm line of division between the types, because
increase of relief or geological differentiation tends to break up the uniform
surfaces of the exterior class: thus the valleys of Moonbi Ranges (Tamworth)
are interior features, with granite floors surrounded by metamorphic ridges, and
some valleys of the Glen Innes region, in basalts, have a similar form. On the
whole, however, the types are well differentiated, and the smaller interior features
are localized. Two sub-types may be recognized, according to the nature of
bounding highlands.

(i) Ridge and Plain.—This division is of great importance in the life of
the State, as the plains furnish easy routes, and are highly productive. They
are interrupted and partially enclosed at least by high ridges, which form up
to 25% of the landscape. The ridge crests in any one district usually lie in
a plane 800 to 2,500 feet above the lower features, and are of an undulating
character. The slopes are steep, and are often rocky and forested, with angular
values up to 60°, and rarely below 20°. The form depends on the nature of the
local rock, but as the ridges are generally determined by the grain of the
country and major joints, they tend to have straight edges and rectangular
patterns. They rise steeply from the plains with only narrow foothill slopes,
generally not exceeding a mile in width, and appear like steep coasts or islands
rising from the sea. On the other hand, the enclosed plains are gentle and
smooth, with slopes varying from 0° to 0° 30’ in any direction, and the streams
usually flow in very slight depressions hardly below the general level. An
exception to this latter rule is found on the southern side of Liverpool Range,
where tributaries of Goulburn River pass to level valleys 500 feet below the
basalt-sandstone plain, but mostly the flood plains of the streams are simply
parts of the valley floor, which does not vary appreciably between the bounding
ridges. Thus the valley of Conadilly River includes such an alluvial plain
20 miles in width.

This leads to another characteristic of the plains, namely, their alluvials.
These are universal, and consist of an upper horizon of fine sand or silt, usually
5 to 10 feet in thickness, and a lower one of rounded pebbles of general thickness
3 to 6 feet where exposed, but in many localities the modern streams are still
flowing over the old pebbles. The process of channel cutting is still going on,
so the plains are exposed to two major actions—sheet flood erosion, and
channelling. The principal occurrences of ridge and plain features are on the
western slopes of New England (Liverpool Plains); the valleys of the Peel and
Conadilly Rivers; the southern and south-west sides of Liverpool Range, leading
to Mudgee and Rylstone districts; Goulburn; the Shoalhaven Valley; the Lake
George-Canberra district; the headwater country of the Murrumbidgee; the basins

444 COASTAL TABLELANDS AND STREAMS OF NEW SOUTH WALES,

of the Eucumbene-Snowy valley, and the upper parts of the Murray (Swampy
Plain) valley. In the examples to the north of the Blue Mountains, the bounding
ridges are generally horizontal lavas or sandstones: to the south, they consist
of long, straight, narrow ridges, and of hill masses in the case of local basins of
the head Murrumbidgee and Eucumbene-Snowy.

(ii) Plateau and Plain.—This type of scenery is almost confined to the
margins of the Triassic sandstones of the Hunter-Sydney region. The higher
bounding features are plateaus, partly of sub-horizontal sandstones and partly
of older rocks, and the enclosed areas are undulating plains of some relief,
generally up to 500 or 600 feet. The two surfaces are separated by cliffs,
frequently precipitous, and are sharply differentiated; the characteristic break
of slope between the bounding scarp and the lower plain obtains, and in addition
there is another major break between the scarp and the upper plateau surface.
It is usual to find the enclosed features divided into steps or terraces: there
are series of such levels along the upper Cudgegong, Colo and Cox Rivers, the
slopes from one to another being slightly roughened in the first case, and
trenched by deep gorges in the third, while the Colo terraces are separated by
unbroken cliffs. As opposed to these, the valley of the Hunter is close to sea-level,
and has alluvial bottom plains in the narrower valleys above Singleton; below
that town, the wider expanse of the valley is hilly and undulating, although
there is also an expansion of the flood plains.

The development of limited interior features may be observed in Burragorang
Valley (lower Wollondilly), where the valley floor attains a maximum width of
two miles; half of which is alluviated. At Newton Boyd and Araluen, valleys
of triangular plan are developed mainly in granites above major stream junctions,
with sides from two to five miles long. In a similar class, perhaps, is the limited
plain along the coast which is enclosed by high plateau on one side, and the
ocean on the other. This occurs between Bellingen and Hawkesbury Rivers, and
from the Illawarra Coast south of Sydney to the Victorian border: it is partly
composed of alluvial or shore line deposits, and ends sharply against the eastern
scarp of the plateau mass, with very limited extensions along the main rivers.

Plateau Scarps and EHEdges.—The main plateau scarps run north and south
on either side of the highlands (Text-fig. 2). To the east, their front is well
defined and continuous, but to the west the edge marked for the highlands is
more arbitrary, owing to the gradual disintegration of the slopes. There is,
however, a broken line of hills marking the eastern limit of the inland plains,
or the western plateau margin, and a rather less continuous fall from the compact
tableland to the nearest of the flanking valleys within the limits of the country
of high relief. Thus the fall to the Murray tributaries in the vicinity of Mt.
Kosciusko is 4,000 to 5,000 feet, that to Bathurst and associated plains from the
Blue Mountains is 2,000 feet, and that from Moonbi Ranges (S.W. of New England)
to the Peel Valley is 3,000 feet. In each case the plan of the edge is complex,
and conforms to the principal drainage lines by sending branches along the
main water partings. These slopes or scarps are not to be confused with the
ridge features of the plateau surfaces, which follow the grain of the country,
and are meridionally disposed. The granite ridges of New England and the
granite and quartzite ridges of the Goulburn-Kosciusko region are typical of the
latter, and are an integral part of the plateau surface: the immediate boundaries
cf the tableland, however, are features or surfaces distinct from either the plateau

BY I A. CRAFT.

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Text-fig. 2.—Relief of the Main Divide and highland edges. The
solid columns point inwards towards the heights of the Main Divide
Gniddle columns), and show the elevation of plateau edges above their
surroundings. The hollow columns towards the west show the elevation
of the local bases above sea-level, but such a value on the eastern side
is too small to be depicted. The difference between the outer columns
(including the hollow) and those of the Main Divide shows the altitude
represented by gentle regional slopes, while the hollow columns show
the vertical distances through which the inland slopes fall to sea-level.
This gives an idea of the relative importance of the main stages in the
rise to the Main Divide.

446 COASTAL TABLELANDS AND STREAMS OF NEW SOUTH WALES,

or lower plains, and are consistently falling slopes whose average value is of the
order of 20°.

From this it will be seen that the slopes east and west of the Main Divide
are broadly divisible into two sections: an upper slope of the summit or crest
plane whose angular value rarely exceeds 2°, and a steep lower slope delimiting
the main plateau blocks, whose value depends on local circumstances, and may
approach 45° in small areas. Northerly or southerly falls are less easy to define
on account of the meridional elongations of ridges, which have irregular crests
and abrupt terminations, as in the case of the extremities of Gourock Range,
but the fall in high or low points of the landscape is of the same order as the
gentler part of the transverse fall noted above, with breaks due to erosional
scarps. Among the latter may be included the northern fall to Darling Downs
(Queensland), the local scarp to the south of Guyra Plain, the southerly fall
of New England to the Peel Valley, the northerly fall of the Blue Mountains
to Mudgee-Cassilis, and the southerly fall of that mass to Lake George Plains,
near Goulburn. These features vary in direction, but all serve to break the
continuity of slope along the Main Divide. In addition to these, there are slopes
associated with the peripheral depressions; these will be noted separately.

Growth of the Plateau.

Turning now to the evolution of the features which have been outlined, one
has to consider two factors—erosion and tectonic action. The history of erosion
can be deciphered because important elements of ancient landscapes have been
preserved by flows of basalt and other lavas. This allows of the reconstruction
of the pre-basalt landscape, and gives a measure of subsequent erosion, and the
nature and amount of uplift. The information thus gained can be used, in turn,
to define the role of peneplanation in the formation of highland surfaces, and to
help elucidate stream history.

Physiographic Meaning of the Basalts.

The Tertiary eruptive rocks of the plateau may be referred to loosely as
“basalts”, after the predominating member of the group. In vertical range they
are found from sea-level to an altitude of 5,700 feet, and in areal distribution
at intervals over the whole plateau, with major and minor concentrations in New
England and Monaro respectively. The flows may be divided into three groups
according to topographical relations, namely: those of the coastal plain; those
which conform to the present surface, and those which are anomalous with
respect to modern conditions. These will be noted in order.

Basalts of the coastal plain.—So far as is known, these are confined to the
coast south of the Shoalhaven River (Nowra). They occur within 300 feet of
sea-level on a limited plain of variable width cut in the edge of the plateau
mass, and are thus newer than the canyons of those slopes, and newer than
the uplifting of that margin of the plateau. Highland basalts of the nearby
region apparently pre-date the canyons, and the coastal examples are quoted
separately to place them outside generalizations relating to the plateau surface.

Conformable to existing topography.—This class includes the greater part
of the areal distribution. In it the basalts occur in valleys which are still
occupied by streams, or the surface they have covered slopes in sympathy with
modern stream basins, with the greatest elevation of lava on or near the major
divides. Thus the basalts of the Monaro region covered a surface whose topography
was generally similar to that of the present Eucumbene, Murrumbidgee and

BY F. A. ORAFT. 447

Snowy Rivers, and exceptionally high points are near the divides, or cover
ancient monadnocks (e.g., Tabletop, 5,580 feet). In the case of the Macleay basin,
New England, the base of the basalt falls from the divides towards the main
stream junctions, and again the high points are mainly built up around ancient
heights, or along persistent divides. In other places, such as Vegetable Creek,
Emmaville, the flows have been separated from their immediate surroundings by
later erosion, but they still lie much below the high points of the landscape at
the stream heads.

Unconformable to existing topography.—The basalts of the Central Tablelands,
between the Hunter Valley and Goulburn, occur largely as isolated heights on the
surface of the tableland and the crests of divides, with limited extensions into
high-level valleys. This is especially the case to the north of a line drawn
westward from Sydney, but further south the isolated cappings give place to
the greater flows of the Taralga-Goulburn district which form the highest points
of that region, and include the Main Divide for a distance of 30 miles.

Relief shown by Tertiary Basalts.

From this approximate division it will be seen that the basalts of the
Central Tableland occur as high features of the scenery, and vary from a little
above to a little below the highest points in respective districts. On the other
hand, those of New England and Monaro-Kosciusko were ejected on a surface oi
much higher relief, so the first conception obtained of pre-basalt conditions is
one of relatively high northern and southern country separated by a lower middle
stretch.

In estimating this past relief, as in any general discussion, the base of the
basalt is referred to. In some instances this base lies on stream deposits in
channels of definite grade. Outstanding examples are Vegetable Creek (New
England) (Text-fig. 3), tributaries of Shoalhaven River, and the Kiandra flow

Text-fig. 3—Talwegs of streams in the Emmaville district, from data collected by
T. W. EH. David. “A” is the ancient basalt-filled channel of Vegetable Creek, preserved
as far downstream as “E”’, with subsidiary basalt-filled channels near the main line
shown by small crosses. “V’’ is the modern grade of Vegetable Creek, and ‘“‘B” that of
Beardy River. For the coincidence of ancient and modern talwegs on the uplands, see
IDYEINAKel (GLEXS7, 105 Be) )s

(south), which occupy channels showing a normal longitudinal profile. In
each case, similar features have been reproduced in detail by modern streams,
and the immediate relief involved is of the order of 1,000 feet. Extending such a
line of argument to Monaro from district to district, it has been found that post-
basaltic erosion above the modern altitude of 2,500 feet has been devoted to the
re-development of earlier features and stream profiles in detail, thus giving a
maximum pre-basalt relief of 3,500 to 4,500 feet above the lowest basalts of the
region, with no probability of great differentiation of altitude during uplift (Craft,

448 COASTAL TABLELANDS AND STREAMS OF NEW SOUTH WALES,

1933). This is a conservative estimate, as the major pre-basalt features of the
neighbouring Shoalhaven Valley are 500 feet lower still, again without great
probability of later change.

Conditions in New England are best inferred from a study of the Macleay
River (Text-fig. 4). Its upper drainage is almost perfectly enclosed by a circle
of radius 35 miles and centre at the junction of Apsley River with the main

le tg ava
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Le

HASTINGS R.

:

SCALE OF MILES
(a) 10 20 20
eS

Text-fig. 4.—Macleay River basin, with basalt cross-hatched or
black, and major waterfalls shown by double lines. The circles are
centred at the junction of Apsley River with the main stream, and have
radii of 22 and 35 miles respectively.

stream, and large tributaries flow inwards towards this centre. Within it (i.e.,
west of the latitude of Bellbrook) 70% of the enclosing divide is of basalt, but
in each portion other rocks rise at intervals to the crest of the basalts, or a little
higher. From this rimming watershed, 4,000 to 5,000 feet above sea-level, there
is a centripetal fall, of the order of 800 feet per 15 miles measured in the general
direction of the streams, and a line of cataracts is reached at 3,000 to 3,300 feet
altitude: the disposition of the basalts is in sympathy with this fall, and the
various terraces or levels at 3,200, 3,500 and 4,000 feet have been largely re-exposed
by the stripping of their basalt covering, with the lowest flows and gravels
remaining towards the stream centre.

Further east, the basalts of Dorrigo and the Hastings ridges generally have
a base about 2,000 feet above sea-level, although it is as low as 1,600 feet to the
south-east of Dorrigo. Comparing this with the upper Macleay basin, the difference
in basaltic altitude is of the order of 1,000 feet, and the centre of the circle
referred to above is 40 miles further west than the town of Dorrigo. Now if the

BY F. A. CRAFT. 449

bases of the basalts in the two localities be compared, and an allowance made for
the pre-basalt fall between them, it is unlikely that the two have been differentiated
by post-voleanic uplift to a greater extent than 500 feet. As the modern course
of the Macleay from the centre of the circle to Kempsey, 16 miles. west of the
coast, is 109 miles, this estimate might allow a comparatively low grade for the
pre-basalt stream. The conclusion is that, irrespective of the absolute ages of
the Macleay and Dorrigo basalts, which may be similar, the amount of later
warping between the Main Divide and the coastal belt in this locality does
not involve a greater vertical displacement than 500 feet. The existing relief
above 2,000 feet is thus essentially older than the basalts.

This conclusion is supported to the north-west of the Macleay, where the
Vegetable Creek basalts mapped by David (1887) lie in a normal channel of
erosion between 2,000 and 3,500 feet altitude. Modern streams have developed
similar profiles to the lower limit (Text-fig. 3), so the probability of considerable
warping on this part of the western slopes is small. In this connection it is
interesting to recall that Andrews (1904) placed the major extrusions of the
region in the first part of the “canyon period’, and both he and David (1887)
quoted instances where canyon streams towards the western limits of these
highlands have not yet cut through basalt-filled channels, the recorded instances
being at a modern altitude below 2,000 feet.

Turning now to the Central Tableland, between the Hunter Valley and
Goulburn, essentially different conditions are met. As a general rule, the isolated
drifts and basalts occur in valleys not more than 500 feet in depth, and some
flows on the northerly divides of the Colo are based on isolated points equivalent
to the highest level of the nearby sandstone surface. The base of the flows in
various districts indicates variable local relief: this has a minimum value of
500 feet towards the north in the Colo basin, a rather similar value in the central
portion about Jenolan, and in the southern about Taralga, and a value of 800
feet in the south-east between Moss Vale and the middle Shoalhaven; half of
the latter figure seems to be due to uplift immediately preceding the basalts.
Arguing solely from this, a general relief of 500 feet may be ascribed to the Blue
Mountains portion of the pre-basalt surface, and a possible altitude of 1,000 feet
above sea-level, with such ancient residuals as Mt. Lambie rising almost to
2,000 feet. There is another factor in addition to those mentioned, as the base oi
the basalt varies greatly from district to district: characteristic altitudes are
3,500 feet (Colo divide), 3,000 feet (Grose basin), 4,000 feet (Shooter’s Hill),
3,800 to 3,000 feet (Mt. Werong to Abercrombie River), 2,700 feet (Taralga), 2,000
to 2,400 feet (eastern Wollondilly, Craft, 1928, p. 642). Now the surface on which
these flows rest has been deformed to the extent indicated by these figures since
the deposition of the Triassic rocks, and the question arises: Was this deformation
earlier or later than the basalts? Up to the present, the evidence has been
construed to gain the answer, “later”, and it would be difficult to arrive at any
other conclusion when the relationships of the canyons intersecting the basalts
of the Wollondilly, Shoalhaven and Grose River districts are studied. If this
answer also applies to the more northerly section, there has been tilting or
faulting towards the low country of the Hunter-Castlereagh districts, but the
possibility of greater earlier elevation in that area has yet to be investigated.

In conclusion of this section it may be stated that, at the time of basaltic
extrusion, the Central Plateau or Blue Mountains formed a low region between

450 COASTAL TABLELANDS AND STREAMS OF NEW SOUTH WALES,

the higher masses. Its general elevation probably did not exceed 1,000 feet, with
a maximum of 2,000 feet, whilst the higher mass of New England rose to 3,500
feet, and the Monaro-Kosciusko region to 4,500 or 5,000 feet as a maximum.
There appears to have been little subsequent deformation in the higher regions,
put the Central Plateau may have been warped considerably.

Changes of Elevation shown by Basalts.

The changes deduced from basalt occurrences depend on the presence of
underlying terrestrial stream deposits, and the consequent assurance that the
land surface affected was above sea-level in basaltic times. It has been suggested
that the presence of the stream gravels indicates widespread subsidence (e.g.,
Andrews, 1910c), but existing streams have deposited gravels at similar grades,
and the channel gradient, arrangement, and other local features of various deposits
make the possibility of general subsidence very remote. Changes involved are thus
due to movements of uplift, except on the littoral, where there is evidence of
both uplift and subsidence with respect to present sea-level, movement in either
direction being of the order of 200 feet. For the plateau surfaces, the changes
of elevation inferred are based on this discrepancy between pre-basaltic profiles
and those of the present day; the eastern side is used in preference to the
western, as it is based on sea-level, while the western base-level on the edge
of the plateau is 500 to 600 feet higher. There is no way of telling how this
latter has varied within a relevant period, as any change of slope due to
uplift has been too slight to allow of the development of distinctive new features
on the inland plains.

Examining the plateaus in detail, it is found that there are three major
areas where the basalt-flows come near sea-level, namely, Brisbane and Richmond
River (3850 and 0 feet altitude respectively), the Hunter-Castlereagh area (700
feet near Merriwa), and the south coastal plain between Nowra and Eden (up
to 300 feet). In the first example, the total movement is small; in the second,
uplift is limited to the order of 500 feet, which is also the depth of gullies below
basalt-filled channels, and the third example post-dates the canyons, and sets a
local limit to most recent uplift of the order of 300 feet. Elsewhere, canyons
have been eroded below the basalt lines to the depth of 2,000 feet in the Clarence,
Macleay and Hastings basins of the North Coast, in the Blue Mountains, and in
the southern tableland, especially in the case of Shoalhaven River. To the west
of the Main Divide, similar gorges rarely exceed 1,000 feet in depth except in
the fall to Murray River, in the edge of the higher Kosciusko mass. As the
higher features approach the coast and are thus exposed freely to erosion from
base-level, it may be granted that 2,000 feet represents the general magnitude
of post-basaltic uplift in the main plateau masses quoted, and there are inter-
mediate and bordering areas where it has been much smaller, namely, Darling
Downs, the Richmond and lower Clarence basins, the Hunter-Cassilis-Castlereagh
area, the Sydney basin and the lower Clyde. All of these may be considered as
relatively stationary areas.

These considerations also disclose the main parts affected by warping, which
comprise the surroundings of the Richmond-Clarence, Sydney and Clyde River
areas, the northern and western fall of New England, the northern and southern
slopes of the Blue Mountains, and the slopes on either side of the Shoalhaven-
Monaro-Kosciusko plateau. This is already well recognized on the evidence of
tilted rocks or inclined planes of erosion, but it must be emphasized that the

BY F. A. CRAFT. 451
oe

tilting involved has been only a portion of the gentler regional slopes referred
to earlier, whose total rarely exceeds 1°, and the steeper tableland edges are
separate erosional features. Where basalts occur on the inclined surfaces of
the main plateaus in the warped zones, as on the northern slope of New
England (Queensland) to the Darling Downs and the fall of the Blue Mountains
south of Jenolan, it is assumed that they have shared the general warping.
Where they cover channels of normal stream profile, especially where these
have been re-developed in more modern times, as on the slopes at Vegetable Creek,
the Macleay basin, Shoalhaven River and Kiandra, the general fall of the surface
(i.e., the whole general angular value up to 2°) is of erosional origin. Reid’s
sections of the northerly fall of the basalts from New England do not conflict
with the details of this interpretation, but if it should be shown that those
basalts occupy unwarped channels of normal profile, it will be necessary to grant
a pre-basaltic relief for northern New England approximating to that of the
present day, as he contends (1926, pp. 304-305), instead of allowing later uplift
of 2,000 feet to complete the existing relief in that area.

Age of Peneplanation.

Having given an indication of pre-basalt relief and the extent of later
movements of uplift, it remains to be seen whether an earlier form of the
landscape can be deduced. The only relevant evidence appears to be offered by
the relationships between the sub-horizontal Mesozoic or earlier rocks, and the
older masses which they flank. The most complete reduction appears to have
taken place in the Blue Mountains, where the surface of unfolded Mesozoic
or Permian rocks gives place to planes cut in granites or older folded strata with
very little change of altitude in the passage. That this relationship is ancient
is shown by the line of hard residuals, or monadnocks, which extends over a
distance of 150 miles between the Mudgee and Goulburn districts. These points
rise from 300 to 600 feet above the newer rocks on their flanks, and were
residuals at the close of the Triassic period, indicating that the older rocks
had been planed down to approximately their present surface at that time.

In the Shoalhaven-Monaro region there is evidence that the upper surface
of the Triassic-Permian strata is continuous with a peneplain in the older bounding
rocks. Different action by combined uplift and erosion may have given the
essentials of the existing landscape above the (modern) height of 3,000 feet by
the end of the Mesozoic era, while that above the modern altitude of 2,000
feet was developed before the period of Tertiary basalts (Craft, 1932, 1933). New
England is still more difficult to discuss, as the intrusion of granites and
concomitant uplift appear to have survived into the Mesozoic. The central mass
is flanked by Mesozoic rocks which are very rarely above 2,000 feet in altitude;
where the passage from newer to older strata is not marked by wide valleys—
such as the Hunter and Peel—there is a sharp rise to the older mass, which thus
appears as a low dome partly enclosed by the more recent surface. It is impossible
to say precisely whether this area was ever reduced to form an approximately
horizontal plane, but something of the kind may have taken place, as the course
of newer movements has been to elevate the central mass, while neighbouring
areas have had their persistent basin form renewed or accentuated.

Summing up, it may be concluded that the most complete peneplanation
recorded in the history of existing land forms is in the region of the Blue
Mountains, where monadnocks are comparatively insignificant points. The New

452 COASTAL TABLELANDS AND STREAMS OF NEW SOUTH WALES,
7

England and Monaro-Kosciusko regions have been domed gently, with a con-
siderable proportion of the movement being post- and pre-Mesozoic respectively.
The highlands thus formed have been subjected to continuous invasion, and
incomplete peneplains have been eroded; these are shallow basins of erosion in
the north and great terraces in the south, which lead to higher cores of gently
convex aspect in each case. Present erosion from east and west is tending to
impose a sinusoidal curve in section from east to west throughout the length
of the highlands.

STREAMS OF THE PLATEAU.

Classification.—On the basis of present knowledge, a stream classification can
be made by reference to profiles and canyon relationships. Three major types
may be recognized, namely, (i) one in which a profile of equilibrium exists,
without notable canyon sections even in the headwater trace; (ii) one in which
a profile of equilibrium has been attained, with appreciable canyon sections; and
(iii) one in which there is a torrent section, and juvenile canyons apart from
the headwater tract, the torrent section usually occurring between graded upper
and lower courses (Text-fig. 5). These may be taken in order.

ee an io |

| | |. GBrishane
| | |
| ]

Namoi

| | Castlerea Bh

Gc | Hun ter

| | Lachlan

Macquarie

Murrumbidgee
| | Snowy

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| OPEL fawkesbury
| ———4
E igees : A Barwon
| PAZ [ Clarence

Ly Ly Macleay

Re
: Shoahaven
LLY
Miles Vertical Sc a/c 000 reese V:H = 33.

lo 100 200 300 00

Text-fig. 5—Stream Profiles (talwegs) of representative streams. Shading indicates
canyon sections, and the letters stand for the following river names: P. = Peel; C. =
Conadilly; T. = Talbragar (to Macquarie); G. = Goulburn; Cx. = Cox; N. = Nepean;
B = Beardy; T.C. = Tooloom Creek; Na. = Nymboida; M. = Mann; A. = Apsley; Ga. =
Guyra; D. = Deua; A.C. = Araluen Creek. The principal lengths are by courtesy of
the Lands Department.

BY F. A. CRAFT. 453

(i) Hxamples: Condamine, Lockyer’s Creek, Brisbane River, Conadilly, Peel,
Castlereagh, Talbragar, Goulburn-Hunter.

These have talwegs with slight upper concavity, and almost the whole of
each course is heavily alluviated, the maximum width of flats for a single
stream being 16 miles in the valley of Conadilly River above Gunnedah. The
banks of the Condamine, Brisbane and Hunter are steep in places where they
pass through or near hard rocks, and local hills rise some hundreds of feet
above them. However, the general topography is that of wide, shallow valleys
or interior plains, with smooth surfaces favouring the operation of sheet floods.
Such conditions are found almost to the heads; thus the Condamine rises in
flat valleys dominated by scattered buttes, Lockyer’s Creek in the basalt scarp of
Darling Downs, and the Conadilly in the northern scarp of Liverpool Range,
but in each case the flow is directly on to plains. The upper Peel and the Hunter
occupy trough-like valleys with rounded and conical hills respectively rising
above them, while the Goulburn Valley has steep sides and a more pronounced
grade for part of its course, although it can hardly be included in the following
class.

(ii) Hxeamples: (a) Lachlan, Murray; (b) Macquarie, Murrumbidgee, Snowy;
(c) Hawkesbury (Nepean, Cox, Colo, Macdonald).

The first two examples have talwegs rather similar to those of the first
type: the Lachlan has a long, shallow gorge section above Cowra, while the
Murray falls steeply in gorges and narrow valleys to 3,000 feet in depth, and
its upper valleys consist of alternating basins and constrictions of the gorge
sides. Neither stream flows over any considerable area of high plateau, but
each has eliminated major waterfalls from its bed, and has established an
approximate equilibrium, with a slight general tendency towards downcutting.

These streams are not so well known as the following three, Macquarie,
Murrumbidgee and Snowy, which have a generally concave talweg, although the
upper portion of each is distinctly convex, with many irregularities. The upper
course of the Murrumbidgee and (Hucumbene)-Snowy have alternate basins and
constrictions: the basin floors are plain interior features, with considerable
alluviation, from which the streams pass into constrictions without a general
steepening of grades. The constrictions have sides up to 50° in slope, and in parts
the whole space between them is occupied by the river bed. Lower down their
courses, at Burrinjuck in the case of the Murrumbidgee and below Bombala on
the Snowy, the streams flow through gorges, whence they pass to the lower
plain sections. Hach of these rivers has been so well adjusted to local conditions
that in the first 150 miles of its course there has been an almost uniform
downcutting of 100 to 200 feet since the period of basalt flows. The third stream
of the group, Macquarie River, lacks the basins and constrictions towards its
head, and flows in undulating land of varying relief, which includes valleys
1,000-1,400 feet deep in the fall towards the Bathurst Plains. Below that
feature, the river passes into hilly country, with sections of canyons down to
1,000 feet below the local plateau level, whence it flows to the inland plains.

In the Hawkesbury group many topographical features are due to the presence
of jointed sub-horizontal sandstones, which give convex profiles in the gorge
units of the Macdonald, lower Colo, and Nepean basins, and precipitous cliffs
in the broader valleys of the upper Colo, Grose and Cox-Wollondilly Rivers.
These wider upper valleys differ from the basins of the Snowy and Murrumbidgee

K

454 COASTAL TABLELANDS AND STREAMS OF NEW SOUTH WALES,

in being strictly localized, and having wide floors trenched by canyons of varying
depths. With each there is a single major constriction where the dipping
Triassic sandstones are crossed in the passage to the Sydney basin.

(iii) Eavamples: Barwon, Gwydir, Namoi, Clarence, Macleay, Wollondilly and
Shoalhaven Rivers.

This type differs from the preceding in having greater lengths of canyons,
and a torrent tract including major rapids and waterfalls. The headwater
section has a concave talweg, and the river grade below the steep middle fall
is comparatively gentle, although the streams are usually turbulent, and cross
many bars of hard rock. Approaching the edge of the highlands, the canyons
widen and give place to valleys of decreasing depth, and to plains as the highland
edge is passed.

Considering the examples in detail, it is found that the rivers of New England
west of the Main Divide flow through gentle valleys between plateau ridges
until they come within 2,500 or 2,000 feet of sea-level, when they fall further
into canyons 500 to 1,000 feet in depth, and finally emerge on level valley floors
in piedmont plains about 1,000 feet above sea-level. The eastern streams of New
England have a higher fall line, about the altitude of 3,000 feet; from this
level they fall into canyons to 2,500 feet in depth, whence they pass to the
plains. Thus the Clarence flows into a peripheral depression and has gentle
grades and open valleys in the Mesozoic rocks about Grafton, but the western
tributaries, Timbarra, Mann and Boyd Rivers, flow in great canyons with a
maximum depth of 2,500 feet about Newton Boyd. A feature of the streams in these
canyons, and of the neighbouring Macleay, is the turbulence of the current even
at the lower grades, and the number of rock bars in each channel, although
the presence of stream gravels and drift up to 50 feet above the rivers indicates
a recent period of quieter flow, and presumably of lesser erosion. The headwater
portions of many of the upper Clarence streams are found in a strip of undissected
plateau 20 miles in width; this is reminiscent of the upland basin of the Macleay
which has already been mentioned (section on basaltic relief).

In others of the coastal rivers, the Richmond and Manning especially, more
than half of each course is less than 500 feet above sea-level. They differ from
the Clarence and Macleay in the virtual absence of higher level plateau from
their heads, and in the presence of numerous steep canyons. In the south,
the Clyde, Deua and Tuross are rather similar to these, and complete profiles
cannot be drawn for them at present.

Towards the south, the Wollondilly and Shoalhaven differ from the northern
streams in matters of detail. In both cases the graded upper section consists
mainly of level plains bounded by higher ridges or plateaus, and the lower part
of each is in a precipice-crowned valley or canyon. The streams in the south,
like the Clarence and Macleay in the north, show the headward progression of a
rejuvenating movement by the extension of canyons into uplands, and by sharp
fall lines. On the western side of the Main Divide, similar phenomena are only
seen in certain rivers draining the western slope of New England, and the other
stream courses have a much more ancient aspect, even in the case of the deep
gorge of the Upper Murray (Kosciusko) area, where the steep head falls begin
close to the divides. However, there has been revival here at some time since
the basaltic period, as the Tumut River flows in a gorge cut more than 2,000
feet through basalts and “leads”, and the Macquarie at Bathurst flows more than

BY F. A. CRAFT. 455

500 feet below a similar occurrence. It should be noted that basalts of the Tumut
divide extend southward to the Murray valley at 1,300 feet, so revival does not
necessarily mean uplift in that section.

In addition to the characteristics already mentioned, there are some common
to all of the rivers. Deposits of coarse gravel overlain by finer drift or silt
occur at all altitudes, and on grades as steep as 1:50. They are not common
in narrow canyons where the streams pursue torrential courses, but even here
they are found occasionally up to 100 feet above the present stream beds, and
they occur freely on the gentler gradients of gorges and narrow valleys. Three
stages may be recognized as affecting them: firstly, streams flowed in rock
channels, and transportation of debris tended to exceed accumulation; secondly,
there was a great accumulation of coarse gravels except under very unfavourable
circumstances, ranging from sea-level to the Murray-Kosciusko plateau, especially
on the level plains of interior features; this was succeeded by a deposit of fine
drift and silt. Thirdly, and most recently, there has been a general erosion
of channels in all these deposits, which is still proceeding: on steeper grades
and along more turbulent streams they have been cut through and the old rock
beds re-exposed to attack; on plains, the streams flow partly on them, or are
in the process of cutting channels in level surfaces. This may point to general
hydrographic changes following upon changes in the amount, nature or seasonal
distribution of rainfall. At the present time the rivers are also liable to great
seasonal changes in volume and to floods.

In conclusion of the section, it may be remarked that the three stream
classes adopted are sufficiently representative to comprise all the streams from
the highlands. So far as the plateau is concerned there is no essential difference
between rivers flowing to the Pacific coast, and those flowing inland, and
differences of such details as canyon depths are mainly brought about by the
base level at the western edge of the highlands being 500 or 600 feet above that
of the eastern, which is sea-level. This factor also affects the vertical distance
through which the respective plateau edges fall, though not the value of individual
slopes, and makes the gradients of western streams more uniform than they
would otherwise be. There are, however, no essential differences that would
enable the plateau streams west of the Main Divide to be classed separately
from the eastern; all present an aspect of stability and adjustment to their various
environments, or this adjustment is taking place by regrading following a process
of rejuvenation.

Plan of the Systems.—The general stream arrangement of New South Wales
and relevant portions of adjoining States depends on the long-continued existence
of four major depressions or basins, occupied respectively by the Darling and
Murray Rivers on the west, and by the Brisbane-Clarence and Hunter-Hawkesbury
systems on the east of the Main Divide. The western examples are marked by
the presence of symmetrical river systems, but such a condition is less apparent
on the east, where more restricted areas and relatively greater variety of structure
and elevation impose special local conditions on streams. The river heads in
plateaus do not flow immediately towards the centres of respective basins, or
toward the coast in the case of independent streams, but generally pursue
meridional courses for distances up to 80 miles, measured in a straight line,
before they turn away to flow eastward or westward.

The stream details of the plateaus have been largely determined by the
grain of the country, the presence of specially resistant features, and the persist-

456 COASTAL TABLELANDS AND STREAMS ‘OF NEW SOUTH WALES,

ence of ancient slopes. All three factors operate in the Monaro-Kosciusko region,
where they have determined the restricted meridional courses of the Shoalhaven,
Murrumbidgee, Cotter, Goodradigbee, Tumut and Indi Rivers, and parts of the
Snowy. In New England, streams of the Macintyre, Gwydir and Namoi Rivers
have a general trend to the north-north-west in sympathy with the grain of the
country, and similar causes have influenced the meridional tributaries of the
Clarence.

It might be noted here that the relative unimportance of southward flowing
streams from the southern slopes of New England may be due to the extra
elevation of that edge through the development of the Hunter Boundary fault;
pending agreement on the age of that feature, the matter cannot be discussed
with assurance, as the whole of the high divides immediately north of the
Hunter valley are concerned in the faulting movements.

When the rocks of the landscape are more homogeneous, the streams take
on a Symmetrical character: the Macleay, Hastings and Manning, and the Hunter
tributaries rising about Barrington Tops are conspicuous examples, but there
has been no comparable development south of the lower Shoalhaven, where the
coastal rivers have only obtained a bare foothold on the plateau. As an example
of the intimate relationship between rock structure, slopes and streams, the two
complex eastern basins may be compared in detail (Text-fig. 6).

40

Text-fig. 6.—Locality Maps of the Brisbane-Grafton and Hunter-Sydney areas,
showing the modern extent of Mesozoic rocks, with outliers, in black, to the east of
Nymboida River, and the position of the main streams.

A.
1b

bo

os

4a,

Co

BY F. A.

CRAFT. 457

Comparison of Grafton and Sydney Basins:

Grafton Basin,

Sydney Basin.

i. Topography.

Section—Level

Plateau to 2,000 feet in Mesozoic rocks
—general slopes from N.W. towards Rich-
mond River and Grafton—voleaniec rocks

Northern plateaus,

interior

features, and streams of the first type.
Plateau 1,500 to 2,000 feet—Mesozoic
and unfolded late Palaeozoic—general

slopes from N.W. and N. towards Sydney

of West Moreton (Toowoomba) and basalts of Liverpool Range, Goulburn
Richmond River areas. River and Colo basin.
Faulted north-eastern margin (Brisbane Faulted northern margin (Hunter
River), folding (pre-basalt). Boundary fault).

B. Middle Section—Warped margins leading down to basins.

North slopes to basin from Macpherson
Range.—Sandstones to 2,000 feet—over-
lain by basalts, to 4,000 feet.

Western Margin—Plateau to maximum
of 5,000 feet—sharp rises to west of
Mesozoic rocks in the form of western
valley sides higher than those to the
east.

No equivalent.

Southern Margin—Rise of Mesozoic rocks
to 1,000 feet and continuation of a pene-
plain surface in older rocks.

Central Basin of Richmond-Clarence
Plains and low plateau to altitude of
300 feet—unfolded sediments over 3,700
feet thick below Grafton—Grafton to
Macleay divide = 48 miles.

North slopes from plateau of 1,000 feet
—general depression along Macdonald
river line.
Western

feet, with

margin—Plateau with 4,400
Mesozoic and Kamilaroi rocks
merging with surfaces of older rocks
with no great changes of altitude
between them—Greater part of rise is
by monoclines west of Sydney, and in
Jenolan Plateau.

A series of flat-bottomed valleys
western margin of Mesozoic rocks.

Southern rise of Mesozoics to 2,500 feet
with a surface continuing to older rocks
(as in Shoalhaven Valley).

on

Central Portion—low plateau to 300
feet, with many local plains—unfolded
sediments much more than 3,000 feet
thick below Sydney—Sydney to northern
Shoalhaven divide = 56 miles.

C. Southern Section—Rising plateaus.

Rise southward and westward to Macleay
divides, from 2,000 to 4,500 feet—basalt
flows on divides.

Southward rise continued over Shoalhaven
Basin—depression near coast and_ sur-
face fall to lower Clyde River.

ii. Stream Equivalents.

Brisbane River—near junction of Meso-
zoic with older roeks—associated with
major fault lines—Lockyer’s Creek from
west.

Main Clarence—Nymboida line keeps
close to western extremity of the Meso-
zoic rocks, and to the east of the higher
mass of New England.

Orara River—flows N.W. in sympathy
with dip of Mesozoic rocks and fall of
surface.

Timbarra River flows northward, and
Boyd and Mann systems flow as normal
streams to the main Clarence-Nymboida
line.

Richmond River falls from north towards
the centre of the basin as an indepen-
dent system.

Hunter River—follows close to limits of
Mesozoic rocks—associated with major
fault lines—Goulburn River from west.

Wollondilly and Kowmung Rivers near
western extremity of Mesozoics, and to
east of higher masses of Jenolan
plateau, and its southern extension.

Nepean River—fiows north along general
line of depression, and tributaries flow

N.W. to the depression in the sand-
stones.
Shoalhaven River runs northward

towards the centre of the basin, although
it turns independently to the sea.

Colo and Macdonald Rivers
depression to Hawkesbury.

follow

458 COASTAL TABLELANDS AND STREAMS OF NEW SOUTH WALES,

Summing up, it is found that the topography and streams of these two areas
are similar. In each case, there is a varied fall into a central depressed area,
with a wide valley and a river system on the northern side. Streams flow from
older to newer, sub-horizontal rocks, where their position seems to have been
determined mainly by weak edges where the newer rocks join the older, by the
slopes of the warped sides of the basin, and by certain tectonic features, such as
very gentle synclines. The Brisbane (Marks, 1933, p. 185), Orara, Macdonald and
Nepean have been quoted as examples of this latter, and it is possible that other
streams have a similar origin. Thus Willan (1923, p. 24) suggested this for the
lower (eastward) course of the Hawkesbury, and Carne’s sections (1908) show
the Grose River as flowing in a slight syncline, and the easterly course of
Capertee River (Colo system) below Glen Alice as occupying the line of a
depression 600 feet deep, shown in both the plateau surface or the top and bottom
of the Kamilaroi (Upper) Coal Measures. The writer suggests that the ultimate
stream origins in these cases are to be found in slight roughening of a virtually
undenuded surface during an early phase of the uplift of the Mesozoic rocks,
and to the establishment of other rivers along the marginal lines of weakness:
the latter received drainage from the higher enclosing masses of older rocks.
There appears to be no good evidence for extensive stream rearrangements away
from the general lines of such a primitive course, and the writer believes that
the existing system can be regarded as static over a long period of time,
commencing before the extrusion of Tertiary lavas, although these have doubtless
been responsible for local variations.

CONCLUSION.

1. The progress of physiographic thought and knowledge in the State is
reviewed briefly. Features common to both earlier and later periods are the
idea of an anticlinal Main Divide, the uniformity of structure and process in
Eastern Australia, and the postulation of extensive marginal faulting. Dis-
tinctively modern ideas include those of the peneplain, and the rearrangement of
stream patterns by the interaction of tectonic influences and dynamic stream
action.

2. The topographic features of the New South Wales plateaus may be grouped
as “exterior” or “interior” features, depending on the respective absence or
Eresence of higher enclosing ground in a district, and a sharp break of slope
where its fall meets lower features.

3. The more recent developments of the plateau can be traced by reference
to old land surfaces and stream channels preserved by Tertiary lavas. These
give a measure of pre-eruption relief, which is found to have been of the order
of 3,000 feet in New England, and 4,500 to 5,000 feet in Monaro-Kosciusko region,
with an intermediate zone of low country generally not exceeding 1,000 feet, with
a few higher points. A master peneplanation is of much greater age, and
only parts of the gentle upper slopes of the plateau to the east and west are
due to warping following pre-basalt uplift. The plateau edges are primarily
erosional features.

4. The plateau streams may be included in three classes: the first has a
profile of equilibrium with no canyons; the second has such a profile with sections
of canyon, and the third has a profile of rejuvenation with canyons forming a
large or predominating feature in the highlands. Such characteristics as alluvial
deposits are held in common. | :

BY F. A. CRAFT. 459

5. The stream plan has been determined by the existence of peripheral
depressions. The grain of the country, zones of hardness and ancient slopes
have acted as determinants along the median of the plateau strip, to give stretches
of meridional courses.

6. A comparison is made between the topography and stream positions of
the eastern depressions: the Brisbane River system is equated with the Hunter,
the Clarence-Nymboida with the Wollondilly, the Richmond with the Colo and
Macdonald, the Orara with the Nepean, and the Timbarra, Boyd, and Mann Rivers
with the Shoalhaven.

Viewed as a whole, the region has been subjected to a gentle post-basaltic
uplift of 2,000 feet on the median line of the highlands, with relatively depressed
portions forming Darling Downs and the low country between the Goulburn and
Castlereagh Rivers. The streams are well adjusted to their environment, or
are regrading parts of their channels as a result of uplift mentioned above, and
new lower plains and levels are extending on the margins, forming a new
terrace on either side of the higher terraced plateau mass.

References.

ANDREWS, E. C., 1900.—Report on the Hillgrove Gold Field. Geol. Surv. N.S.W., Min.
Res. No. 8, Sydney.
,1901.—Report on the Kiandra Lead. Geol. Surv. N.S.W., Min. Res. No. 10,
Sydney.
, 1902.— Preliminary Note on the Geology of the Queensland Coast, etc. Proc.
LINN. Soc. N.S.W., xxvii, 146.
, 1903a.—Outline of the Tertiary History of New England. Rec. Geol. Surv.
N.S.W., vii, Pt. 3, 140-216.
, 1903b.—Notes on the Geology of the Blue Mountains and Sydney District.
Proc. LINN. Soc. N.S.W., xxviii, 786.
, 1904a.—Geology of the New England Plateau, Pt. I—Physiography. Rec.
Geol. Surv. N.S.W., vii, Pt. 4, 281-300.
———_ —.,, 1904b.—-Introduction to the Physical Geography of New South Wales. Wm.
Brooks, Sydney.
,1905.—Geology of the New England Plateau, Pt. II—General Geology. Rec.
Geol. Surv. N.S.W., viii, Pt. 2, 108-152.
, 1906.—New Zealand Sound (and Lake) Basins, and the Canyons of Eastern
Australia, etc. Proc. LInN. Soc. N.S.W., xxxi, 499-516.
, 1907.—Geographical Significance of Floods. Proc. LINN. Soc. N.S.W., xxxii,
795-834.
, 1908.—Report on the Drake Gold and Copper Field. Geol. Surv. N.S.W., Min.
Res. No. 12, Sydney.
—, 1910a.—Geographical Unity of Eastern Australia in Late and Post-Tertiary
Time. Journ. Roy. Soc. N.S.W., xliv, 420.
,1910b.—An Excursion to the Yosemite (California). Journ. Roy. Soc. N.S.W.,
xliv, 262.
, 1910c.—Forbes-Parkes Gold Field. Geol. Surv. N.S.W., Min. Res. No. 13,
Sydney.
, 1911.—Erosion and its Significance. Journ. Roy. Soc. N.S.W., xlv, 116.
, 1912.—Notes on a Model of New England. Journ. Roy. Soc. N.S.W., xlvi, 143.
,1914.—New England Tableland. B.A.A.S., Handbook for N.S.W., Sydney.
, 1915.—Geological Survey of the Cargo Gold Field. Geol. Surv. N.S.W., Min.
Res. No. 19, Sydney.
Berry, A., 1825.—Geology of part of the Coast of New South Wales, in Barron Field’s
“Geographical Memoirs on New South Wales”. John Murray, London.
CARNE, J. E., 1908.—Geology and Mineral Resources of the Western Coalfield. Mem.
Geol. Surv. N.S.W., Geology No: 6:7 ~~~
CLARKE, W. B., 1853.—Papers Relative to Geological Surveys, N. S. Wales. Reports
viii and ix, fep., Sydney.
, 1860.—Researches in the Southern Gold Fields of New South Wales. Sydney.
, 1875.—Notes on Deep Sea Soundings. Journ. Roy. Soc. N.S.W., ix, 1-72.

460 COASTAL TABLELANDS AND STREAMS OF NEW SOUTH WALES.

CLARKE, W. B., 1876.—Deep Oceanic Depression off Moreton Bay. Journ. Roy. Soc.
N.S.W., X, 75-82.
CraFt, F. A., 1928.—Physiography of the Wollondilly River Basin. Proc. Linn. Soc.
N.S.W., liii, 618.
, 1932.—Physiography of the Shoalhaven River Valley. Proc. Linn. Soc. N.S.W.,
lvii, 245. vi. Conclusion. (Parts i to v will be found in the same PROCEEDINGS, lvi
and lvii.)
, 1933.—Surface History of Monaro, N.S.W. Proc. LINN. Soc. N.S.W., lviii,
229-244.
Davip, T. W. E., 1887.—Geology of the Vegetable Creek Tin-Mining Field. Mem. Geol.
Surv. N.S.W., Geology No. 1.
, 1896a.—Anniversary Address. Journ. Roy. Soc. N.S.W., xxx, 33.
, 1896b.—See Etheridge, R., David, T. W. E., and Grimshaw, J. W.
————,, 1902.—An Important Geological Fault at Kurrajong Heights. Journ. Roy. Soc.
N.S.W., xxxvi, 359.
, 1932.—Explanatory Notes to accompany a new Geological Map of the Com-
monwealth of Australia. Sydney.
Davin, T. W. E., and ETHERIDGE, R., 1890.—Raised Beaches of the Hunter River Delta.
Rec. Geol. Surv. N.S.W., ii, Pt. 2, 67.
EXTHERIDGE, R., Davin, T. W. E., and GrimsHAaw, J. W., 1896.—Occurrence of a Sub-
merged Forest, with Remains of the Dugong. Journ. Roy. Soc. N.S.W., Xxx, 158.
HEDLEY, C., 1910.—Submarine Slope of New South Wales. Proc. LINN. Soc. N.S.W.,
3O:0-ay, Ys
,1911.—Study of Marginal Drainage. Proc. LINN. Soc. N.S.W., xxxvi, 13.
JENSEN, H. I., 1906.—Preliminary Note on the Geological History of the Warrumbungle
Mountains. Proc. LINN. Soc. N.S.W., xxxi, 228.
, 1907.—Geology of the Nandewar Mountains. Proc. LINN. Soc. N.S.W., xxxii,
842.
Marks, E. O., 1933.—Some Observations on the Physiography of the Brisbane River and
neighbouring Watersheds. Proc. Roy. Soc. Q’land., xliv, No. 9, 132.
MURCHISON, R., 1845.—Cited by Clarke, W. B., 1860 (q.v.).
Rew, J. H., 1926.—Tertiary Unity or Disunity in Queensland. Rept. A.A.A.S., Vol. 17,
pp. 300-312, 1924.
SUssmiitcH, C. A., 1909.—Notes on the Physiography of the Southern Tableland of
N.S.W. Journ. Roy. Soc. N.S.W., xliii, 331.
, 1911.—Geology of New South Wales. Govt. Printer, Sydney.
, 1914.—Physical Geography of New South Wales, ete. B.4.A.8., Handbook
for N.S.W., Sydney, 486-505.
, 1923.—Presidential Address. Journ. Roy. Soc. N.S.W., Ivii.
SUssmMILonH, C. A., and JENSEN, H. I., 1909.—-Geology of the Canobolas Mountains. Proc.
LInn. Soc. N.S.W., xxxiv, 157.
Taytor, T. G., 1907.—Lake George Senkungsfeld. Proc. Linn. Soc. N.S.W., xxxii, 325.
, 1910.—Physiography of the Proposed Federal Territory at Canberra. Com-
monwealth Bureau of Meteorology, Bulletin No. 6.
————.,, 1911.—Physiography of Eastern Australia. Commonwealth Bureau of
Meteorology, Bulletin No. 8.
,1923a—Warped Littoral around Sydney. Journ. Roy. Soc. N.S.W., lvii, 58.
, 1923b.—Blue (Mountain) Plateau. Pan-Pacifiec Sci. Cong., Guide Book to Blue
Mountains, 17-29.
THompson, A. M., 1869.—Notes on the Geology of the Country around Goulburn. Trans.
Roy. Soc. N.S.W., iii, 57.
WILKINSON, C. S., 1882.—Notes on the Geology of New South Wales. Dept. of Mines,
Sydney.
WILLAN, T. L., 1923.—Geology of the Sydney District. Pan-Pacific Sci. Cong., Guide
Book to the Sydney District, 24.

AUSTRALIAN HESPERIIDAE. IV.
Notes AND Descriptions or New Forms.
By G. A. WATERHOUSE, D.Sc., B.E., F.E.S.

[Read 29th November, 1933.]

Brigadier W. H. Evans has sent me further critical notes as the result of his
investigations on this family at the British Museum. It is fortunate that the
British Museum has now acquired the specimens of this family that belonged to
the late H. Fruhstorfer, so Brigadier Evans has been able to examine most of
Fruhstorfer’s types and to supplement the inadequate published descriptions.
Fruhstorfer seems to have had very little appreciation of generic distinctions,
and although he states (/ris, xxiv, 1910, p. 60) that he based his classification on
that of Watson, and Elwes and Edwards, his use of genera often differs from
these authors. Fruhstorfer makes little, if any, use of antennal characters and
male brands, which distinguish species that are otherwise similar in general
appearance. A number of Mabille’s types are now also in the British Museum.

Mr. T. G. Campbell has recently sent a fine collection of skippers from
Brock’s Creek, North Australia. These specimens are now in the Australian
Museum and with the Division of Economic Entomology, Canberra. Besides the
species mentioned later, the following new and important records are in this
collection: Toxidia sexguttata sela Waterhouse, 1932 (February); Neohesperilla
senta Miskin, 1891 (January to March); Neohesperilla xanthomera Meyrick and
Lower, 1902 (February, March); Ocybadistes hypomeloma vaga Waterhouse, 1932
(February and June).

This part adds to and corrects part ii in These Procrepinas for 1932, where a
list of the chief references is given (p. 238). The types of all the new races are
in the Australian Museum, Sydney, and typical specimens are in the British
Museum.

TRAPEZITES IACCHOIDES Waterhouse.

During September, 1932, this rare species was searched for at Como, N.S.W.,
without success, but five males were caught during October of that year. In
September, 1933, it was on the wing, showing that the spring of 1932 was a late
one.

A new locality has been found near Hawkesbury Lookout, Springwood, where
the species was not uncommon during September and October, 1933. Females were
exceedingly rare, not more than four being seen. Most of the males were
caught in an area of about one hundred square yards.

TRAPEZITES PHIGALIOIDES Waterhouse.

Males of this species were not uncommon on the Blue Mts., at Mt. Victoria
and Katoomba, in December, 1932. Two females only were caught at Mt. York.
Males have also been caught at Killara in October, 1932, and 1933.

L

462 AUSTRALIAN HESPERIIDAE. iV,

HESPERILLA CHAOSTOLA Meyrick.

The holotype of this species is a male from Blackheath, caught in November.
It is a rare species in New South Wales, as I have until recently had only six
males and two females, all from the Blue Mts.

In December, 1932, at Mt. Victoria, I found feeding on Gahnia Sieberi several
nearly full-grown larvae which were unknown to me. The larvae of Hesp. donnysa
were plentiful on the same plant and on two occasions I found both types of
larvae on the same clump of Gahnia. An attempt to get the Gahnia to grow in
Sydney failed. The larvae were left at Mt. Victoria and visited each month until
May, 1933. -I found that they grew very slowly and had not pupated in May.
Most had pupated in September and six pupae were brought to Sydney. In October
a pair emerged and proved to be this species.

The life cycle is of extreme interest. All larvae found in December, 1932,
were in the last instar, except one, which was just about to enter the last instar.
In January, 1933, a further search was made and several very small larvae were
found, which were not beyond the second instar. These small larvae were from
eges laid in October or November, 1932, whilst the large larvae were undoubtedly
from eggs laid in October or November, 1931.

The larva is of the usual Hesperilla type, but is more highly coloured than
that of the other species. The segment behind the head is bright red and the
head is covered with longish white hairs. During the daytime the larva lives
in a shelter made by joining the tips of the leaves of the Gahnia together and
enters from below and lives head downwards. It pupates in the same shelter,
also head downwards, fastened by the tail, but without a silken pad. This
method of pupation is quite different from the other species of the genus, but is
similar to that of Motasingha atralba. The pupa is nearly black and the pupal
cap is somewhat similar to that of Hesp. donnysa, but the corrugated platform is
not raised so much.

The life cycle of two years of this species is remarkable, more especially as
the closely allied species, Hesp. donnysa, feeding on the same plant in the same
locality, has a life cycle of one year.

Hesp. chaostola must be rare on the Blue Mts., as some thousands of plants
were searched by myself and three schoolboy friends and only six pupae were
found. At the same time many dozen larvae of Hesp. donnysa were found. It is
one of the earliest Hesperids on the wing, appearing about the middle of October.

Now that perfect specimens are available from the Blue Mts., I am able
to confirm a previous opinion that Victorian specimens constitute a new race,
as follows.

HESPERILLA CHAOSTOLA CHARES, N. Subsp.

Upperside not nearly so dark as in the typical race, spots slightly larger,
especially those in 4 and 5 of forewing, and more heavily scaled, often an extra
spot in 2 (not in holotype); usually indistinct patches of scales parallel to termen
of hindwing, cilia of hindwing paler.

Underside: Apex of forewing and hindwing lilac-grey and not markedly dark
purple as in the typical race.

The greatest difference is shown by the underside, which is very dark, even
in poor specimens of the typical race, and very much paler in chares.

Holotype male, Beaconsfield, Vict., 17 Oct., 1904 (Fig. 691 in Butterflies of
Australia, Waterhouse and Lyell, 1914); allotype female, Beaconsfield, 26 Oct.,

BY G. A. WATERHOUSE. 463

1904 (Fig. 690, loc. cit.); paratype males and females, Beaconsfield, 17 Oct. to
7 Nov., 1904 (all caught by Dr. W. IE. Drake) in Australian Museum. I have
specimens from Ringwood, Vict., and Mr. A. N. Burns caught two females in the
Grampians, Vict., in Nov., 1931.

Tasmanian specimens probably constitute another race, but I have only one
male for comparison.

MOTASINGHA DIRPHIA DEA, n. Subsp.

Since my previous paper I have caught several more specimens of M. dirphia
on the Blue Mountains. These are smaller, with smaller and paler markings, and
are considerably darker in both sexes than the race dilata Waterhouse, 1932,
found at the sea coast near Sydney.

The holotype male from Blackheath (November) has three distinct subapical
dots, a cell spot and a spot in area 3 on the upperside of the forewing. On the
underside of the hindwing it has the large central white spot ringed with black
and one discal white dot. Some males from Blackheath have only one or two
subapical dots, others have a spot in area 2 of the forewing. I have no male
with three white discal dots on the underside of the hindwing; one usually is
present in area 6 and often dark dots in areas 2 and 3.

The allotype female (Blackheath, December) has three distinct subapical
dots, cell spot and spots in areas la, 2 and 8 of the forewing, but not extended
along the veins as in dilata. The underside of the hindwing has the central white
spot ringed with black and three discal white dots. Other females have an obscure
cell spot on the upperside of the hindwing and two of the three discal spots on
the underside of the hindwing represented by black dots.

This race has been taken at Blackheath from November to February; Mt.
Victoria, December and January; Katoomba, November; Wentworth Falls,
November. Specimens from Berowra and Killara are nearer this race than dilata.

TARACTROCERA Butler.
Catalogue Fabrician Diurnal Lepidoptera, p. 279, 1869.
This genus is distinguished by the spatulate and excavate antennal clubs with
very short or entirely obsolete apiculi.

TARACTROCERA DOLON Plotz.

Apaustus dolon Plotz, Stett. Ent. Zeit., 1884, Dp. 165.—T. dolon, Waterhouse
and Lyell, Butt. Aust., 1914, p. 201, figs. 876-7.

The description of Plotz is poor and his unpublished coloured figure rather
misleading. The name dolon was a MS. one given by Herrich-Schaeffer. It is not
possible to say from where the type came, but very probably from near
Rockhampton. Brigadier Evans believes that the species figured in the Butterflies
of Australia is the one described by Plotz, but I incline to the opinion that if the
type is ever discovered, it will be found to be a race of Ocybadistes walkeri Heron,
1894; however, the name can remain for the present. In eastern Australia the
butterfly figured in the Butterflies of Australia occurs from Brisbane to the Cairns
district. The following race from North Australia presents considerable
differences.

TARACTROCERA DOLON DIOMEDES, 1. subsp.
Larger and brighter and with broader markings than specimens from eastern
Queensland. In appearance somewhat like a small 7. ina Waterhouse, 1932, but
the subapicals of the forewing are not divided by darker veins and the male has

464 AUSTRALIAN HESPERIIDAE. iv,

raised scales over veins la, 2, and sometimes below 3, whereas the male of T. ina
has no sex brand. In both sexes the undersides are a very much brighter orange
than in dolon. In the male the spot in area 6 on the hindwing is very variable in
size.

I have had this race in my collection from Darwin (September) for some
years, and it is also in the British Museum from the same locality. Mr. Campbell
has sent a series from Brock’s Creek, North Australia, caught from January to
April.

TARACTROCERA INA Waterhouse.

Proc. Linn. Soc. N.S.W., 1932, p. 228.

This is a rare species, as Mr. Campbell was only able to secure four specimens
at Brock’s Creek in February and March. These agree with the type in the South
Australian Museum, from Darwin. Further material both from North Australia
and Queensland is necessary to determine if the specimens I listed from Queens-
land in 1932 are a distinct race or not. However, a small series described below
presents certain differences.

TARACTROCERA INA IOLA, N. subsp.

In this race the orange bands of the upperside are darker and broader than
in the typical race. It is also a smaller insect. Six males and three females were
caught on Hayman Is., Whitsunday Group, by Mr. F. A. McNeill, of the Australian
Museum staff, in January. The male has no sex brand on the forewing. The
female is somewhat similar in shape to the female of 7’. anisomorpha Lower, 1911,
but the subapicals of the forewing are divided by the darker veins and the under-
side is not so uniformly suffused with orange-yellow.

TARACTROCERA ANISOMORPHA Lower.

Bibla anisomorpha Lower, Trans. Roy. Soc. S. Aust., 1911, p. 146. —T. bavius
anisomorpha, Waterhouse, Proc. Linn. Soc. N.S.W., 1932, p. 229.

Brigadier Evans has examined the type of 7. bavius Mabille from Timor, now
in the British Museum, and finds it is a race of T. nigrolimbata, so T. anisomorpha
will rank as a full species. Mr. Campbeli has taken it at Brock’s Creek in January,
February and April.

TARACTROCERA ILIA Waterhouse.

T. udraka ilia Waterhouse, Proc. Linn. Soc. N.S.W., 1932, p. 229.

The type of 7. udraka Fruhstorfer, 1910, could not be found in his collection,
now at the British Museum, and Brigadier Evans considers that, as Fruhstorfer’s
description agrees much better with Ocybadistes ardea Bethune-Baker, 1906, wdraka
is best considered as identical with ardea. T. ilia will then rank as a full species.

OcYBADISTES Heron.

Ann. Mag. Nat. Hist., 1894, xiv, p. 105.—Padraona, Waterhouse (nec Moore),
Proc. Linn. Soc. N.S.W., 1932, p. 230.

Brigadier Evans considers that Padraona is best restricted to Oriental species;
eastern allied forms have more or less flattened antennal clubs and a different
type of male genitalia and should be placed in the genus Ocybadistes, type
O. walkeri Heron, 1894.

OCYBADISTES FLAVOVITTATA Latreille.
Hesperia flavovittata Latreille, Encyclopédie Méthodique, ix, p. 768, 1824.—
Padraona hespera Waterhouse, Proc. Linn. Soc. N.S.W., 1932, p. 233.

BY G. A. WATERHOUSE. 465

Brigadier Evans has made a critical examination of the specimen mentioned
‘in my previous paper on page 232. This specimen, now in the British Museum,
was originally in the Boisduval collection and bears Latreille’s label flavovittata.

It is without doubt one of the specimens mentioned when Latreille wrote his
description. It is a female and belongs to the species with a narrow sex brand
in the male. Therefore my name hespera falls as a direct synonym. This being
so, the synonymy of my previous paper requires considerable alteration. The
earliest name with any certainty for the species with the broad sex brand in the
male is O. walkeri Heron, 1894, typically from Damma Is. Both species vary in
a similar manner in Australia, so the following names are supplied for them.
O. flavovittata so far has only been found in Australia, but O. walkeri extends
beyond Australian limits.

OCYBADISTES FLAVOVITTATA VESTA Waterhouse.
Padraona hespera vesta Waterhouse, Proc. LINN. Soc. N.S.W., 1932, p. 234.
This is the race found typically at Port Darwin. It is small and bright. Mr.
Campbell found it commonly at Brock’s Creek from January to April and in June.
Specimens from North Queensland constitute the next race.

OCYBADISTES FLAVOVITTATA CERES, ND. Subsp.

Padraona hespera vesta Waterhouse, Proc. Linn. Soc. N.S.W., 1932, p. 234
(in part).

This name is introduced for specimens caught by myself at Kuranda, Qld.,
in June. It occupies an intermediate position between the race vesta and typical
flavovittata from Sydney. They are larger, but not so bright as vesta and smaller
and duller than flavovittata. They correspond to the race sonia of walkeri found
also in North Queensland. I have this race from Mackay to the Cairns district.

OCYBADISTES FLAVOVITTATA FLAVOVITTATA Latreille.
Padraona hespera hespera Waterhouse, Proc. Linn. Soc. N.S.W., 1932, p. 233.
This race typically is from Sydney, but is found from South Queensland
through New South Wales and Tasmania. It is apparently absent from Victoria.

OCYBADISTES WALKERI Heron.

Ann. Mag. Nat. Hist., 1894, xiv, p. 106.

The type locality assigned to this species is Damma Is. Port Darwin is
given as a second locality. Brigadier Evans informs me that these latter
specimens constitute a distinct race, mentioned below. The Australian races
vary in the same way as those of O. flavovittata, with an additional race,
hypochlora Lower, 1911, from South Australia. These races are supplied with
names as follows.

OcYBADISTES WALKERI SOTHIS, N. subsp.

Padraona flavovittata flavovittata, Waterhouse, Proc. Linn. Soc. N.S.W., 1932,
19s ZB

This name replaces that of flavovittata as the result of finding the female
specimen in the Oberthur collection, now in the British Museum, bearing the
label flavovittata. It is the commonest Ocybadistes found at Sydney and has a
broad sex brand in the male and is decidedly greenish on the underside. It has
already been well described. The type series consists of five males and two
females reared from a batch of eggs laid by one female at Sydney in April, 1932.
The butterflies emerged in October with a pupal duration of five weeks. It is

466 AUSTRALIAN HESPERIIDAE, iv,

found at Sydney from August to April and has a longer period of flight and is
commoner than flavovittata. Its range extends from Brisbane throughout coastal
New South Wales. It is also found in Tasmania.

OcYBADISTES WALKERI SONIA, 0. subsp.

This name is introduced for specimens typically from Kuranda. It is some-
what similar to the previous race, is constantly smaller and the underside is not
so conspicuously green. I have it from Kuranda in July, September and October;
also from Cooktown and Mackay. It corresponds to the race ceres of flavovittata.

OCYBADISTES WALKERI OLIVIA, nN. subsp.

Padraona flavovittata walkeri, Waterhouse, Proc. Linn. Soc. N.S.W., 1932,
Die2oo-

This race is very distinct from typical walkeri from Damma Is., having
rounded wings and broad markings. The female before me has only a slight
trace of green colour on the underside. I have it from Port Darwin in
September, and Brock’s Creek in February and April.

TrLicota Moore.

Scudder’s selection of augias Linn. as the genotype of Astycus Hiibner cannot
stand, as the first time it was used by Hubner he included only European species.
It was also used as a genus of beetles about the same time. The species listed
in my previous paper must now be placed under Telicota Moore, genotype
Papilio augias Linn.

TELICOTA KREFFTI ARGILUS, n. subsp.

This race has rounder wings and broader and brighter markings than typical
kreffti, the holotype male of which is in the Australian Museum from Cape
York. It is also slightly smaller and the underside of the hindwing has the
band well developed, whilst in typical kreffti the underside of the hindwing is
almost without markings.

Holotype male and allotype female from Port Darwin without dates. It has
been caught at Port Darwin in May, November and December and is in the
British Museum from Baudin Is.

The only species found so far in this district with which it might be
confounded is the paler and narrower winged 7’. augias argeus.

SOME FURTHER OBSERVATIONS ON THE REARING OF CERATODUS.
By Tuos. L. BAncrort, M.B., Ch.M., C.M.Z.S.
(Communicated by I. M. Mackerras, M.B., B.Sc.)
(Two Text-figures. )

[Read 25th October, 1933.]

These observations are supplementary to my paper read before the Society
on 27th June, 1928 (These PROCEEDINGS, liii, p. 315). It had been shown that
the larval Ceratodus lie out of the water on the mud-banks at intervals during
the day and that, if prevented from doing so (due to the water being too deep),
they die before reaching the age of ten weeks.

The suggestion made to use small vessels, such as enamelled pie-dishes, proved
to be tedious when practised on a large scale, as it entailed a lot of work in
adding water to replace that lost through evaporation. It was also found almost
impossible to keep so small a body of water sufficiently cool in very hot weather.
Subsequently a hatchery was erected which has given the best results hitherto
obtained. Sufficient embryological material was secured to satisfy all the workers
in Europe, one of whom wrote to me: ‘We have all the stages of Ceratodus so
it does not matter now if the animal becomes extinct.”

The hatchery (Text-figs. 1-2) consists of a 2,000-gallon tank, which delivers
river water into the rearing chambers and into the water-jacket which surrounds
them, the overflow being collected in a 1,000-gallon tank, whence it is returned
to the upper tank by means of a No. 38 semi-rotary pump.

The upper tank is made of corrugated galvanized iron. It is circular,
10 feet in diameter by 4 feet high. From the bottom corrugation, a 3-inch delivery
pipe is led off, which connects through a T-piece with two brass stopcocks. One
of these is 4 inch in diameter and feeds into the narrow compartment at the
right-hand end of the breeding chambers, the other is ? inch in diameter and
feeds into the water-jacket.

The breeding chambers are built of 1-inch pine, and consist of a trough 12 feet
3 inches long by 12 inches wide by 9 inches deep, all measurements over all,
divided by wooden partitions into twelve equal chambers, with a small chamber
1 inch in diameter at the right-hand or upper end. All parts must be so put
_together that the whole forms a solid, water-tight job. The right-hand end of
the trough is solid, but all the partitions are provided with slots 8 inches by
1 inch, situated 4 inches above the bottom, and covered on the right-hand or
upper side with perforated zinc. The left-hand or lower end of the trough is
slotted to take an overflow pipe, which is protected by gauze and delivers into
the lower tank. The trough is fitted with a 1-inch cleat which raises the right
hand end and ensures a flow of water through the chambers. A gauze cover,
made in three sections, is provided to protect the open top of the trough.

468 FURTHER OBSERVATIONS ON THE REARING OF CERATODUS,

The water-jacket is of galvanized sheet-iron, 12 feet 9 inches long by 1 foot
6 inches wide by 6 inches deep. The trough is held in place in it by six
galvanized iron straps. The jacket is set dead level and provides a water space
3 inches in diameter all round the trough.

The lower tank is of corrugated galvanized iron and is circular, 7 feet in
diameter by 4 feet high. It is so placed that the overflow from the breeding
chambers delivers into its top. A 13-inch pipe is taken off from the lowest
corrugation. This connects with the pump, which returns the water to the
upper tank.

2
Tos wy.
Text-figures 1-2. Ceratodus hatchery.
1.—Side elevation. 2.—Ground plan. w.j., water jacket; br. t., breeding tank.

A mud-bank was arranged in each of the twelve compartments and a
continuous trickle of water was maintained through the whole system. Twenty
to thirty larval Ceratodus, two weeks old, were placed in each compartment.

Rolled oats, dried egg, Huchytra worms, duck-weed, Spirogyra and other Algae,
also dried liver, were given as food. The fowl eggs and bullock’s liver were
boiled, then grated up and dried in the sun and stored in bottles. A pinch of
the rolled oats rubbed up with a pinch of the dried egg was fed to each compart-
ment twice a week, and a pinch of the liver, some Huchytra worms and Algae
were given about three times a month.

It was observed that the young Ceratodus do not grow evenly, some shooting
ahead whilst others lag behind. At three months old, of twelve preserved at this
date, the largest was 40 mm. and the smallest 23 mm.; the lengths were: 40, 35,
31, 30, 30, 29, 29, 28, 28, 26, 26, 283 mm. At five months the largest of eighteen
was 44 mm. and the smallest 26 mm.; the lengths were: 44, 438, 41, 40, 40, 36, 36,
35, 35, 35, 35, 35, 35, 33, 32, 29, 28, 26 mm. At eight months the largest of twenty-
five was 55 mm. and the smallest 26 mm.; the lengths were: 55, 55, 53, 52, 49, 47,
45, 44, 44, 48, 41, 41, 41, 41, 40, 40, 38, 37, 35, 35, 32, 31, 30, 27, 26 mm. At ten
months the largest of seventeen was 62 mm. and the smallest 40 mm. At twelve
months, of nineteen preserved at this date, the largest was 72 mm. and the
smallest 33 mm.

It would be interesting to know what is the cause of the different rates of
growth. In-breeding might have something to do with it; the robust larvae
may be from parents well mated, whereas the puny larvae may be the progeny
of sisters and brothers. Possibly sex might account for it, yet there is no disparity
in size between the sexes in aged fish, very large specimens up to eighty pounds in
weight of both sexes having been taken. i

BY T. L. BANCROFT. 469

In past work on the embryology of Ceratodus, investigators have mentioned
length of the larvae irrespective of age. This may have led to erroneous con-
clusions. Take for instance one of my three months larvae at 35 mm. and a
35 mm. larva from the five months lot and one of 35 mm. from the eight months
series: these, although of the same length, would show different development.

Of three juveniles preserved when two years and three months old, the lengths
were 181 mm., 153 mm. and 150 mm., and the weights in gms. 40:8, 22-4 and
17:0. Of four, two years and nine months old, the largest was 262 mm. long and
the smallest 175 mm. One preserved when five years old was 16% inches in length
(420 mm.) and weighed 16 oz. (454 gms.).

I am indebted to Mr. E. C. Speering, contractor, for preparing a sketch and
detailed specifications of the hatchery.

TAR

ire

ABSTRACT OF PROCEEDINGS.

ORDINARY MONTHLY MEETING.
29th Marcu, 1933.

Professor A. N. Burkitt, M.B., B.Sc., President, in the Chair.

The President referred to the sudden death of Mr. H. W. Hamilton, which
occurred on Monday evening, 27th March, 1933. Mr. Hamilton had been a member
of the Society since 19381.

The Donations and Exchanges received since the previous Monthly Meeting
(30th November, 1932) amounting to 53 Volumes, 300 Parts or Numbers, 37
Bulletins, 10 Reports and 15 Pamphlets, received from 146 Societies and
Institutions and 5 private donors, were laid upon the table.

PAPERS READ.
1. The May-flies of the Kosciusko Region. i. Introduction and Family
Siphlonuridae. By R. J. Tillyard, M.A., Se.D., D.Sc., F.R.S.
2. The Life-history of Grevillea robusta (Cunn.). By P. Brough, M.A., B.Sc.,
B.Sc.Agr.

ORDINARY MONTHLY MEETING.
26th Apri, 1933.

Professor A. N. Burkitt, M.B., B.Sc., President, in the Chair.

Mr. T. H. Guthrie, Whale Beach; and Mr. Gilbert Wright, Sydney University,
were elected Ordinary Members of the Society.

The President announced that the Council had elected Dr. H. S. H. Wardlaw,
Mr. EH. Cheel, Professor T. G. B. Osborn and Dr. C. Anderson to be Vice-Presidents
tor the Session 1933-34.

The President announced that the Council had elected Dr. G. A. Waterhouse
to be Honorary Treasurer for the Session 1933-34.

A letter was read from Mrs. H. W. Hamilton and son, returning thanks for
sympathy.

The President expressed the congratulations of members to Mr. P. Brough
on attaining his Doctorate of Science in the University of Sydney.

The Donations and Exchanges received since the previous Monthly Meeting
(29th March, 1933) amounting to 14 Volumes, 84 Parts or Numbers, 4 Bulletins,
4 Reports and 15 Pamphlets, received from 54 Societies and Institutions, were
laid upon the table.

PAPERS READ.
1. Some Climatological Aspects of Aridity in their Application to Australia.
By J. Andrews, B.A., and W. H. Maze.
2. Seasonal Incidence and Concentration of Rainfall in Australia. By J.
Andrews, B.A., and W. H. Maze.
N

xXxXX ABSTRACT OF PROCEEDINGS.

8. Notes on Australian Diptera. xxxiii. By J. R. Malloch. (Communicated
by F. H. Taylor.)

4, Revision of Australian Lepidoptera. Oecophoridae. Part ii. By A. J.
Turner, M.D., F.E.S.

5. On the Production of Fertile Hybrids from Crosses between Vulgare and
Khapli Emmer Wheats. By W. L. Waterhouse, D.Sc.Agr.

NOTES AND EXHIBITS.

Mr. T. C. Roughley exhibited specimens of yearling Sea Trout (Salmo trutta)
from the trout hatcheries on the River Plenty, near Hobart, Tasmania. These
fish were covered externally and on the gills by a ciliate infusorian, /chthyoph-
thirius multifiliis, the life history of which was described at the meeting of the
Society on 28th September, 1932. A heavy mortality of Sea and Rainbow Trout
occurred in the River Plenty hatcheries during February, 1933, and, although
much of the trouble is thought locally to be due to the stirring up of the bottom
where considerable quantities of unconsumed food had accumulated and putrefied,
there is little doubt that the parasites caused the deaths of many of the fish. The
liberation in Tasmania of this parasite, which has been introduced from abroad
on aquarium fishes, is fraught with great danger to the freshwater fishes of that
State.

Mr. T. C. Roughley also exhibited specimens of an Isopod crustacean known
as a Sea Louse and identified by H. M. Hale, the Director of the South Australian
Museum, as Cirolana corpulenta Hale. These were obtained from sharks and rays
caught in nets by Mr. Norman Caldwell off Stockton (Newcastle) Beach on
24th and 25th December, 1932. The sharks captured (135 in number) comprised
Whalers (Carcharhinus macrurus), Grey Nurse (Carcharias arenarius), Tiger
Sharks (Galeocerdo arcticus), and White Sharks (Carcharodon carcharias), and
all were badly affected with the parasites, which were literally eating them alive.
The sharks, which were captured for their commercial products, smelt very
offensively when opened up, the attacks being general throughout the internal
organs, the gills, liver, stomach, intestine, and flesh being badly infested. The
fiesh had turned a brownish colour, the intestines were perforated, and the livers,
which in some cases contained hundreds of the parasites, were greatly reduced
in bulk and were very ragged. The whole of the commercial products were
ruined, and all the sharks had to be discarded. Previously, on 14th December,
1932, several sharks captured at Port Stephens were found infected with the
parasites, but not nearly so badly as at Newcastle.

Mr. G. P. Whitley exhibited a right-hand male and a left-hand female of
the Four-eyed Fish (Anableps dowei Gill) from Honduras and made remarks
upon their structure. He also showed the cranium of a Catfish (Neoarius australis
Gtinther), from Moreton Bay, Queensland, the ventral surface of which resembled
a crucifix,

ORDINARY MONTHLY MEETING.
31st May, 1933.
Professor A. N. Burkitt, M.B., B.Sc., President, in the Chair.
The President expressed the congratulations of members to Sir Edgeworth

David on the award of the degree of Doctor of Science by the University of
Sydney.

ABSTRACT OF PROCEEDINGS. xxxi

The President also offered the congratulations of members to Mr. Alan Burges,
Miss Joyce Vickery and Miss Enid Edmonds on their attaining the degree of
Master of Science of the University of Sydney; also to Miss Germaine Joplin,
B.Se., on the award of a Junior Fellowship by the International Federation of
University Women.

The President announced that Dr. Walkom had received an invitation from
the Geological Society of America to attend the 16th International Geological
Congress at Washington, 22nd—29th July, 1933, and had been granted the necessary
leave of absence by the Council to enable him to accept.

The Donations and Exchanges received since the previous Monthly Meeting
(26th April, 1933) amounting to 9 Volumes, 90 Parts or Numbers, 2 Bulletins,
6 Reports and 9 Pamphlets, received from 55 Societies and Institutions and 2
private donors, were laid upon the table.

PAPERS READ.
1. The Petrology of the Hartley District. ii. The Metamorphosed Gabbros
and Associated Hybrid and Contaminated Rocks. By Germaine A. Joplin, B.Sc.
2. Australian Coleoptera. Notes and New Species. No. 8. By H. J. Carter,
B.A., F.E.S.
3. Corynebacteria as an Important Group of Soil Micro-organisms. By H. L.
Jensen, Macleay Bacteriologist to the Society.

NOTES AND EXHIBITS.
Mrs. C. A. Messmer exhibited specimens of a new species of Pterostylis from
Fitzroy Falls.

ORDINARY MONTHLY MEETING.
28th JUNE, 1933.

Professor A. N. Burkitt, M.B., B.Sc., President, in the Chair.

Mr. Horace J. Willings, B.A., Sydney, was elected an Ordinary Member of the
Society.

Letters were read from Mr. Alan Burges, Miss Enid Edmonds, Miss Germaine
Joplin and Miss Joyce Vickery, thanking members for congratulations.

The President announced that Dr. Walkom left on Thursday last, 22nd instant,
to attend the 16th International Geological Congress, to be held at Washington,
22nd-29th July, and is expected to return to Sydney about 9th September next.

The Donations and Exchanges received since the previous Monthly Meeting
(31st May, 1933) amounting to 21 Volumes, 172 Parts or Numbers, 4 Bulletins,
3 Reports and § Pamphlets, received from 70 Societies and Institutions and 1
private donor, were laid upon the table.

PAPERS READ.
1. Notes on New South Wales and Queensland Orchids. By Rev. H. M. R.
Rupp, B.A.
2. The Marine Plankton of the Coastal Waters of New South Wales. Part i.
By Professor W. J. Dakin, D.Sc., F.Z.S., and A. Colefax, B.Sc.

NOTES AND EXHIBITS.

Professor W. J. Dakin exhibited two spider crabs, male and ripe female,
from Corner Inlet, Victoria, where a remarkable invasion has occurred and
considerable damage to fishermen’s nets has resulted therefrom. The specimens

XXxli ABSTRACT OF PROCEEDINGS.

are apparently Leptomithrax spinulosus, although the larger ones are of greater
size than those recorded from various parts of the Australian coast. The Victorian
Chief Inspector of Fisheries stated that specimens up to five feet across had been
obtained. According to the fishermen the crabs first appeared in such large
numbers as to be a pest about three years ago, and each year since has been
worse, the invasion occurring in the winter months. Enormous numbers of this
spider crab have, however, been noted on South Australian shores during winters
many years ago, and there is evidence that invasions occurred on the Tasmanian
coast forty years ago. The unusual migration of exceptionally large numbers of
Leptomithraxz spinulosus into shallow coastal waters may be added to the list
of interesting cases (mostly of terrestrial animals) of extreme fluctuations in the
density of animal populations.

Miss Germaine Joplin exhibited various plants from the Hartley District,
grown on a very siliceous felsite and similar to the flora supported by the
Hawkesbury Sandstone.

Dr. G. A. Waterhouse exhibited two species of butterflies from Hayman Is.,
Queensland, caught by Mr. F. A. McNeill during two visits this year. Taractrocera
ind, a recently described species, was caught in January with three specimens of
Padraona tanus nihana. In May the former species was absent, but a long series
of the latter species was obtained.

ORDINARY MONTHLY MEBRTING.
26th JuLy, 1933.

Professor A. N. Burkitt, M.B., B.Sc., President, in the Chair.

The President referred to the death of Mr. Clive Lord, Curator of the
Tasmanian Museum, Hobart, and Local Hon. Secretary of the Australian and
New Zealand Association for the Advancement of Science.

The Donations and Exchanges received since the previous Monthly Meeting
(28th June, 1933) amounting to 14 Volumes, 137 Parts or Numbers, 1 Bulletin,
5 Reports and 9 Pamphlets, received from 74 Societies and Institutions and 1
private donor, were laid upon the table.

PAPERS READ.

1. Vegetative Reproduction in Drosera peltata and D. auriculata. By Joyce W.
Vickery, B.Sc.

2. The Surface History of Monaro, N.S.W. By F. A. Craft, B.Sc., Linnean
Macleay Fellow of the Society in Geography.

3. On the Distribution, Habitat and Reproductive Habits of certain Huropean
and Australian Snakes and Lizards with particular Regard to their Adoption of
Viviparity. By H. Claire Weekes, D.Sc., Linnean Macleay Fellow of the Society
in Zoology.

NOTES AND EXHIBITS.

Rev. H. M. R. Rupp exhibited a large collection of sketches in crayon and

black-and-white of Australian orchids.

ORDINARY MONTHLY MEETING.
30th Aucust, 1933.
Professor A. N. Burkitt, M.B., B.Sc., President, in the Chair.
Mrs. Edith Coleman, Blackburn, Victoria, was elected an Ordinary Member
of the Society.

ABSTRACT OF PROCEEDINGS. xxxili

The Donations and Exchanges received since the previous Monthly Meeting
(26th July, 1933) amounting to 58 Volumes, 165 Parts or Numbers, 7 Bulletins,
9 Reports and 4 Pamphlets, received from 87 Societies and Institutions, were
laid upon the table.

PAPERS READ.

1. A new Genus and Species of Australian Proctotrypidae. By Alan P. Dodd.

2. The Scientific Name of the Commercial Oyster of New South Wales. By
Tom Iredale and T. C. Roughley.

3. The Life History of the Australian Oyster (Ostrea commercialis). By
T. C. Roughley, B.Sc., F.R.Z.S.

ORDINARY MONTHLY MEETING.
27th SEPTEMBER, 1933.

Mr. E. Cheel, Vice-President, in the Chair.

The Chairman announced that the Council is prepared to receive applications
for four Linnean Macleay Fellowships tenable for one year from ist March, 1934,
from qualified candidates. Applications should be lodged with the Secretary, who
will afford all necessary information to intending candidates, not later than
Wednesday, ist November, 1933.

The Donations and Exchanges received since the previous Monthly Meeting
(30th August, 1933) amounting to 5 Volumes, 63 Parts or Numbers, 3 Bulletins,
3 Reports and 19 Pamphlets, received from 44 Societies and Institutions, were
laid upon the table.

PAPERS READ.
1. The Coccidae of the Casuarinas. By W. W. Froggatt, F.L.S.
2. Useful Coccinellidae found on the Comboyne Plateau. By E. C. Chisholm,
M.B., Ch.M.
3. The Geology of the South Coast of New South Wales, with Special
Reference to the Origin and Relationships of the Igneous Rocks. By Ida A.
Brown, D.Sc.

NOTES AND EXHIBITS.
Mr. E. Cheel gave a short account of the Fifth Pacific Science Congress.
Mr. E. G. Jacobs exhibited (i) a flower spike of the “Common Sun Orchid”,
Thelymitra ixioides Swartz, in which each of the two lowest flowers has three
distinct labella, all three of the same size, and arising from the same region in
front of the central column. All the other flowers on the spike are normal. The
specimen was collected on south shore of Middle Harbour. (ii) Three flowers of
Caladenia alba R.Br., one of which has three labella, and each of the other two
has two labella. In all cases the labella are of normal size, and have the usual
specific characters. The specimen was collected near Fuller’s Bridge, Chatswood.
Mr. G. P. Whitley exhibited a Long-finned Eel (Anguilla reinhardtii
Steindacher) from Misima Island, Papua. This eel is common in fresh water in
Hastern Australia but has not hitherto been recorded from New Guinea.

ORDINARY MONTHLY MEETING.
25th OcToBErR, 1933.
Professor A. N. Burkitt, M.B., B.Sc., President, in the Chair.
Messrs. L. A. Judge, Hornsby, and W. H. Maze, Burwood, were elected
Ordinary Members of the Society.

XXXIV ABSTRACT OF PROCEEDINGS.

Candidates for Linnean Macleay Fellowships, 1934-35, were reminded that
Wednesday next, lst November, is the last day for receiving applications.

The President announced that an invitation had been received from the
Australian National Research Council for members to attend the first David
Lecture to be delivered by Professor EH. W. Skeats, of the University of Melbourne,
at Science House, Gloucester Street, Sydney, on Friday, 17th November, 1933, at
8 p.m. The title of the Lecture will be ‘Some Founders of Australian Geology”’.

The President announced that the North Queensland Naturalists’ Club
proposes to hold a camp at Low Island on the Great Barrier Reef from 30th
March to ist April, 1934.

The Donations and Exchanges received since the previous Monthly Meeting
(27th September, 1933) amounting to 4 Volumes, 80 Parts or Numbers, 1 Bulletin,
1 Report and 3 Pamphlets, received from 50 Societies and Institutions and 3
private donors, were laid upon the table.

PAPERS READ.

1. Notes on the Australian Species of the Family Paussidae. By the late
T. G. Sloane. (Arranged for publication by H. J. Carter, B.A.)

2. The Genus Pterostylis (Orchidaceae). A New Scheme of Classification,
with Notes on the Distribution of the Australian Species. By Rev. H. M. R.
Rupp, B.A.

3. Some Further Observations on the Rearing of Ceratodus. By Thos. L.
Bancroft, M.B., Ch.M., C.M.Z.S. (Communicated by I. M. Mackerras, M.B., Ch.M.)

4. An Investigation of the Sooty Moulds of New South Wales. i. Historical
and Introductory Account. By Lilian Fraser, M.Sc., Linnean Macleay Fellow
of the Society in Botany.

5. Miscellaneous Notes on Australian Diptera. i. By G. H. Hardy.

NOTES AND EXHIBITS.

Dr. A. B. Walkom gave a short account of the Sixteenth International
Geological Congress.

Rev. H. M. R. Rupp exhibited specimens of Cryptanthemis Slateri recently
obtained at Bullahdelah.

Dr. J. McLuckie contributed the following preliminary note on Cryptanthemis
Slateri Rupp.

This plant is a holosaprophytic type, subterranean in habit and devoid of
chlorophyll.

It is apparently rootless, and represents a sympodially branching rhizome,
each segment of which terminates in a capitulum of small flowers. The lower
part of the rhizome has numerous hairs, which have a multicellular base, and
the mycorrhizic fungus was seen in the bases of these hairs.

The mycorrhizic fungus is a typical Orchidaceous type, very similar to that
of Dipodium punctatum in habit, branching, and method of penetrating from
cell to cell. It is extremely like a Rhizoctonia in structure and habit, but is
distinctly coarser in texture than that of Dipodium.

In the outer part of the rhizome, the mycelium is seen aS numerous separate
branching hyphae or in the form of pelotons, very similar to those which appear
on Rhizoctonia in pure culture: the infected cells of this zone show the typical
Pilzwirthzellen phase of the mycorrhiza. Within this zone, a well-defined zone,
usually one layer broad, is occupied by the Verdauungzellen stage, where the

ABSTRACT OL PROCEEDINGS. XXXV

fungal hyphae are undergoing digestion. The Verdauungzellen cells show very
considerable enlargement, and in them all stages of fungal digestion are proceeding.
Very few dextrin granules were seen, but in the Verdauungzellen and adjacent
cells numerous oily globules and a tremendous amount of proteid matter were
seen.

From my preliminary examination, it appears that a fatty material takes
the place of the dextrin so abundant in Dipodiuwm, while there is a surprising
amount of a proteid material in the cells where digestion is proceeding.

The young seedling of this plant appears to be rootless, but rhizoidal structures
are present, and infection by a fungus, probably the mycorrhizal-forming one, is
present. It seems that infection takes place early, and is probably obligatory
for development.

A more complete account of this interesting holosaprophytic orchid will
be published shortly.

Mr. G. P. Whitley, on behalf of Dr. T. L. Bancroft, exhibited a series of
specimens of Ceratodus.

Dr. G. A. Waterhouse exhibited pupae of Heteronympha cordace reared from
eggs laid by captive females caught at Blackheath in January, 1933. All the
species of this genus of butterflies have now been bred and all the species of
Satyridae known to occur in New South Wales. He also showed a coloured
drawing of a pupa found at Mt. Kosciusko in December, 1921. This pupa did
not emerge, and until this year it was not known to what species it belonged.
The pupa is suspended by the posterior end and is bright yellow-green with
jet-black markings. It is more handsome than any satyrid pupa yet known from
Australia. Coloured drawings, one green and one brown, of the very variable
larvae were also shown. The head is hard, hairy, with two slight lateral
projections which are more prominent in the early instars.

ORDINARY MONTHLY MEETING.
29th NoveMseErR, 1933.

Dr. C. Anderson, Vice-President, in the Chair.

Miss Florence R. W. Burton, B.A., Haberfield, was elected an Ordinary Member
of the Society.

The Chairman referred to the death of Dr. T. L. Bancroft on 12th November,
at Wallaville, Queensland.

The Chairman announced that the Council had reappointed Mr. F. A. Craft,
B.Sc., Miss Lilian Fraser, M.Se., and Dr. I. V. Newman, M.Sc., to Linnean Macleay
Fellowships in Geography, Botany and Botany respectively for one year from
1st March, 1934; and had appointed Mr. N. Alan Burges, M.Sc., to a Linnean
Macleay Fellowship in Botany for a period of one year from 1st March, 1934.

The Donations and Exchanges received since the previous Monthly Meeting
(25th October, 1933) amounting to 22 Volumes, 191 Parts or Numbers, 16 Bulletins,
5 Reports and 18 Pamphlets, received from 77 Societies and Institutions, were
laid upon the table.

PAPERS READ.

1. The Coastal Tablelands and Streams of New South Wales. By F. A. Craft,

B.Se., Linnean Macleay Fellow of the Society in Geography.

XXXVi ABSTRACT OF PROCEEDINGS.

2. The Mycetozoa of New South Wales. By Lilian Fraser, M.Sc., Linnean
Macleay Fellow of the Society in Botany.

3. A New Species of Pterostylis. By (Mrs.) Pearl R. Messmer.

4. Australian Hesperiidae. iv. Notes and Descriptions of New Forms. By
G. A. Waterhouse, D.Sc., B.E., F.E.S.

NOTES AND EXHIBITS.

Dr. G. A. Waterhouse exhibited several of the new races described in his
paper on Hesperiidae together with typical specimens of other races for
comparison. He also showed the early stages of Hesperilla chaostola with a life
cycle of two years and of Hesperilla donnysa with a life cycle of one year.

Rev. H. M. R. Rupp exhibited (i) a plant of Cleisostoma Nugentii Bailey in
flower from Proserpine, North Queensland; (ii) enlarged photographs of
Cryptanthemis Slateri Rupp; and (iii) a series of coloured sketches of Australian
orchids.

Professor W. J. Dakin and Mr. A. N. Colefax exhibited a plankton catch,
about which they contributed the following note: On Sunday, 5th November, 1933,
whilst obtaining the usual sea water samples at a station some four miles east
of North Head (Port Jackson), the surface of the sea was noticed to be much
agitated by myriads of small fish, probably anchovies, on which the seagulls
were feeding with great eagerness. An attempt was made to capture some of
these fish in the large coarse-meshed plankton net, but met with no success.
However, even during the few moments that the net was in the water, many large
(from the planktonic viewpoint) Crustacea were found to be entangled in the
net material, and a short haul was made immediately. This haul proved to be of
an interesting and unusual nature. Apart from the abundant and beautifully
preserved Ctenophores, there were numerous large Crustacea. In point of
numbers the most common form was the larva of the Euphausid Nyctiphanes
australis, which, incidentally, has been taken at approximately this time in past
years. The large Copepod, Calanus brevicornis, was also very abundant, as was
the Copepod Sapphirina gemma. The most unusual feature of the haul, however,
was the capture for the first time of really large numbers of Hyperiid Amphipods
which, overseas, have proved to be so important in the food-chains of commercially
valuable fish. The principal genus amongst this haul of Amphipods was
Themistella, and it is interesting to note that this genus was not taken by the
“Challenger” when she was in these waters.

It is possible that the presence of the large Crustacea is to be associated with
the occurrence of the rather smaller ones (Copepods), and also that there is a
close connection between the activity of the fish and the dense swarm of pelagic
Crustacea.

Miss L. Fraser exhibited specimens of Dictydium rutilum, a species of
Mycetozoa newly described by Miss G. Lister (Journ. Bot., August, 1933).

DONATIONS AND EXCHANGES.
Received during the period 1st December, 1932, to 29th November, 1933.
(From the respective Societies, etc., unless otherwise mentioned. )

ABERYSTWYTH.—Welsh Plant Breeding Station, University College of Wales.
Bulletin, Series H, No. 13 (1932); “The Welsh Journal of Agriculture”, ix
@933))r:

ADELAIDE.—Department of Mines: Geological Survey of South Australia. Annual
Report of the Director of Mines and Government Geologist for 1931 (1932);
Mining Review for the Half-Years ended June 30th, 1932 (No. 56) (1932) and
December 31st, 1932 (No. 57) (1933).—Field Naturalists’ Section of the Royal
Society of South Australia and South Australian Aquarium Society. “The
South Australian Naturalist”, xiv, 1-4 (1932-1933).—Public Library, Museum
and Art Gallery of South Australia. Records of the South Australian Museum,
v, 1 (1933). —Royal Society of South Australia. Transactions and Proceedings,
Ivi (1932) —South Australian Ornithological Association. “The South Aus-
tralian Ornithologist”, xii, 1-4 (1933).—University of Adelaide. “The Aus-
tralian Journal of Experimental Biology and Medical Science’, x, 4 (T.p. & c.)
(1932); xi, 1-3 (1933)—Woods and Forests Department. Annual Report for
Year ended June 30th, 1932 (1933).

ALBANY.—New York State Library, University of the State of New York. New
York State Museum Bulletin, Nos. 290, 291, 294, 298; New York State Museum
Handbook, 13-14 (1932-1933).

ALcER.—Société d'Histoire Naturelle de VAfrique du Nord. Bulletin, xxiii, 7-9
(T.p. & ec.) (19382); xxiv, 1-6 (1933).—Station d@Aquiculture et de Péche de
Castiglione. Bulletin, 1931, 1-2 (Index) (1932).

AMSTERDAM.—Nederlandsche Entomologische Vereeniging. Entomologische
Berichten, viii, 187-192 (1932-1933); Tijdschrift voor Entomologie, Ixxv, 3-4
and Supplement (T.p. & c¢.) (1932); Ixxvi, 1-3 (1933); Verslagen van de
Vergaderingen der Afdeeling Nederlandsch Oost-Indie van der Nederlandsche
Entomologische Vereeniging, i, 4 (1933).

ANN ARBOR.—University of Michigan. Contributions from the Museum of
Palaeontology, T.p. & c. for Vol. iii; iv, 1-5 (1932); Miscellaneous Publications
of the Museum of Zoology, Nos. 23-24 (1932); Occasional Papers of the Museum
of Zoology, T.p. & ec. for Nos. 216-235 (Vol. x) (1930-1931); Nos. 240-259
(1932-1933); Papers of the Michigan Academy of Science, Arts and Letters,
XVii-xviii, 19382 (1933).

AUCKLAND.—Auckland Institute and Museum. Records, i, 4 (1933).

BALTIMORE.—Johns Hopkins University. Bulletin of the Johns Hopkins Hospital,
Ib, B45 (No, ee 1G) (Gey hb, eG (anaes) on) (GIBB S3))R bbb ales} (GIG BR) 8
University Circular, N.S. 1932, 9-11 (1932); 1933, 1-7 (1933).

XXXVIii DONATIONS AND EXCHANGES.

BANDOENG.—Opsporingsdienst Dienst van den Mijnbouw in Nederlandsch-Indie.
Bulletin of the Netherlands East Indian Volcanological Survey, Nos. 58-60
(T.p. & Index for Nos. 29-60) (1932-1933); 61-62 (1933); Wetenschappelijke
Mededeelingen, Nos. 19, 23-24 (1932-1933); Vulkanologische en Seismologische
Mededeelingen, No. 12 (1933).

BARCELONA.—Academia de Ciencias y Artes de Barcelona. Boletin, vi, 4 (1933);
Memorias, xxii, 28-31 (T.p. & c.) (1932); xxiii, 2-11 (1932-1933); Nomina
del Personal Academico, 1932-1933 (1932).

BASEL.—Naturforschende Gesellschaft. Verhandlungen, xli, 1929-1930 (1931);
xliii, 1931-1932 (1933); Autoren- und Sachregister der Hefte i-x (1834-1852)
der Berichte uber die Verhandlungen und der Bande 1-40 (1852-1929) der
Verhandlungen. Von Dr. C. Walter (1930).

Batavia.—Koninklijke Natuurkundige Vereeniging in Nederlandsch-Indie.—
Natuunkundice tjdschritt) xctiae2 (psc, Cc.) (1932) ise xcilin Tem GL933)r

Brercen.—Bergens Museum. Arbok, 1932, 1-2 (T.p. & c.) (1982-1933); 1933, 1
(1933); Arsberetning, 1931-1932 (1932).

BERKELEY.—University of California. Publications: Botany, T.p. & c. for Vol. xi
(1922-1930); xvii, 3-6 (1933); Entomology, T.p. & c. for Vol. v (1929-1932) ;
vi, 1-6 (1932-1933); Geological Sciences, T.p. & c. for Vol. xx (1932); xxi, 8
(T.p. & c.) (1932); xxii, 3 (1932); xxiii, 1 (1933); Physiology, viii, 2-3 (1932-
1933) ZO OLOL Wa pXXexvas 3-15) GDA eC.) ClI932)) i= xexexaval elie (pander Cs) LOS ais
oo.qnnnl, Gals, (Aas ce ©) (GIGBYRILG GR) S >:0:6-ab:q ales} (108333) 3 dl al) ((al)3333))

BERLIN.—Botanische Garten und Museum. Notizblatt, xi, 107-109 (1932-1933) .—
Zoologische Museum. Mitteilungen, xviii, 2-3 (T.p. & c.) (1932-1933).

BEeRn.—Naturforschende Gesellschaft. Mitteilungen ad. Jahre 1932 (1933);
Verhandlungen, 112-113. Jahresversammlung (1931-1932).

BrrMINcHAM.—Birmingham Natural History and Philosophical Society. List of
Members, 1933, and Annual Report for 1932; Proceedings, xvi, 4 (1933).

BLOEMFONTEIN.—Nasionale Museum. Argeologiese Navorsing, i, 5-7; Paleontologiese
Navorsing, ii, 4-5 (1932).

Botoena.—Laboratorio di Entomologia del R. Istituto Superiore Agrario di Bologna.
Bollettino, v (1932-1933).

Bompay.—Bombay Natural History Society. Journal, T.p. & c. for xxxv, pts. 3-4;
xxxvi, 1-3 (1932-1933) —Haffkine Institute. Report for the Year 1931 (1933).

Bonn.—Naturhistorische Verein der Preussische Rheinlande und Westfalens.
Sitzungsberichte, 1930-1931 (1932); Verhandlungen, Ilxxxvii, 1930-Ixxxviii, 1931
(1931-1932).

Boston.—American Academy of Arts and Sciences. Proceedings, Ixvii, 5-13
(Ger es) NGl93 21933) eixavan= 1-6) CL933)r

BrisBaAnE.—Department of Agriculture. Queensland Agricultural Journal, xxxvili,
6 (T.p. & c.) (1932); xxxix, 1-6 (T.p. & c.) (1933): xl, 1-5 (1933) —Depart-
ment of Mines: Geological Survey of Queensland. “Queensland Govern-
ment Mining Journal’, xxxiii, Oct.-Dec., 1932 (T.p. & c.) (1932); xxxiv,
Jan.-Nov., 1933 (1933).—Great Barrier Reef Committee. Reports, iv, 1 (1933).
—Queensland Museum. Memoirs, x, 3 (1933).—Queensland Naturalists’ Club
and Nature-Lovers’ League. ‘The Queensland Naturalist”, viii, 4-6 (1932-
1933).—Royal Society of Queensland. Proceedings, xliv, 1932 (1933).

DONATIONS AND EXCEIANGES. B 6.9.4 bs

Brno.—Prirodovedecka Fakulta, Masarykovy University. Spisy (Publications)
(Botanical Only), Cis. 170 (1933).

Brookiyn.—Botanical Society of America. “American Journal of Botany”, xix,
8-110) Cl py Gey Gl932ii xx, 1-31(1933)))

Brusse_s.—Académie Royale des Sciences, des Lettres et des Beaux-Arts de
Beigique. Annuaire, 99° Année, 1933 (1933); Bulletin de la Classe des
Sciences, 5° Série, xviii, 1932, 6-12 (T.p. & c.); xix, 1933, 1-4 (1932-1933). —
Musée Royal d'Histoire Naturelle de Belgique. Bulletin, viii, 5, 7, 11-35
(T.p. & c.) (1932); ix, 1-8, 10 (1933); Mémoires, 52 (1932); Mémoires, Hors
Serie (Résultats Scientifiques du Voyage aux Indes Orientales Néerlandaises),
i (1933); ii, 11-12; iii, 12; iv, 3-7; vi, 1 (1932).—Société Entomologique de
Belgique. Bulletin and Annales, Ixxii, 6-12 (Index) (1932-1933); Ixxiii, 1-5
(1933 ).—Société Royale de Botanique de Belgique. Bulletin, Ixv, 1 (1932).—
Société Royale Zoologique de Belgique. Annales, lxii, 1931 (1932).

Bupaprest.—_Wusée National Hongrois. Annales Historico-Naturales, xxvii, 1930-
LO Sie CLIS O=L9Sda)e

Buenos AirESs.—Museo Nacional de Historia Natural. Anales, xxxv (1927-1932).—
Sociedad Argentina de Ciencias Naturales. Revista “Physis”’, xi, 38 (T.p. & c.)
(1932).

BUITENZORG.—Department van Landbouw, Nijverheid, en Handel. Bulletin du
Jardin Botanique, Serie iii, xii, 3-4 (T.p. & c.) (1932); ‘“Treubia’, vii,
Supplement, Livr. 7 (1932).

CaEn.—Société Linnéenne de Normandie. Bulletin, 8° Série, iv, 1931 (1932);
Mémoires, Nouvelle Série, Section Géologique, ii (1932).

CAIRNS.—North Queensland Naturalists’ Club. North Queensland Naturalist, i, 8
@i9sans

CALcuTTA.—Geological Survey of India. Memoirs, lxii, 1 (1933); lxiii, 1 (1933);
Memoirs, Palaeontologia Indica, N.S. xxi, 1 (1933); Records, Ixvi, 1932, 2-4
(T.p. & c.) (19383); Ixvii, 1 (1933).—J/ndian Museum. Memoirs, ix, 6 (1933);
xii, 1-2 (1932); Records, xxxiv, 3-4 (T.p. & c.); xxxv, 1-2 (1932-1933); Report
on the Zoological Survey of India for the Years 1929-1932 (1932).

CAMBRIDGE, Hngland.—Cambridge Philosophical Society. Biological Reviews and
Biological Proceedings, viii, 1-4 (T.p. & c.) (1933).—University of Cambridge.
Abstracts of Dissertations approved for degrees in the University of Cambridge
for Academical Year 1931-32 (1933).

CAMBRIDGE, Mass.—Museum of Comparative Zoology at Harvard College.—Annual
Report of the Director for 1931-1932 (1932); Bulletin, lxxiv, 3-6 (1932-1933);
ixexevae eS (GSS)

CANBERRA.—Commonwealith Bureau of Census and Statistics. Official Year Book,
No. 25, 1932 (1933) —Council for Scientific and Industrial Research: Sixth
Annual Report for the Year ended 30th June, 1932 (1932); Divisions of
Entomology and Plant Industry, Contributions, Nos. 20, 23, 24, 36, 56, 60
@933))2

Canton.—Botanical Institute, College of Agriculture, Sun Yatsen University.
“Sunyatsenia’”’, i, 2-3 (1933).

xl DONATIONS AND EXCHANGES.

Care Town.—Royal Society of South Africa. Transactions, xxi, 1-2 (19382-1933) —
South African Museum. Annals, xvi, 3 (T.p. & c.) (1933); Report for the
Year ended 31st December, 1932 (1933).

Cuicaco.—Field Museum of Natural History. Leaflet, Geology, 13 (1932);
Publications, Botanical Series, viii, 6; xi, 4; Zoological Series, xviii, 10-11
(1932).

CHRISTCHURCH.—Canterbury Museum. Records, T.p. & ec. for iii (1926-1932) ;
dive ot eGL933)i

CLus.—Gradina Botanica. Bulletin, T.p. & c. for ix (1929); xii, 3-4 and Appendix
1 (T.p. & c.) (1932); xiii, Appendix 1 (1933); Contributions Botaniques de
Cluj, i, 1928, 11-21 (T.p. & c.) (1930); ii, 1-4 (1929-1932); Guide de la
Sixiéme Excursion Phytogéographique Internationale, Roumanie, 1931, rédigé
par Al. Borza. Réimpression émendée (1931).

Cormpra.—Universidade de Coimbra: Museu Zoologico. Memorias e Estudios,
Serie i, No. 1, Fasc. 9-13 (1931); Nos. 58-61, 64 (1932).

CoLtp Spring Harsor.—Department of Genetics: Carnegie Institution of Washington.
Annual Report of the Director, 1931-1932; Report of the President of the
Carnegie Institution for Year ending October 31, 1932 (extracted from Year
Book, Carnegie Institution of Washington, No. 31, 1931-32) (1932).

CoLomBo.—Colombo Museum. Spolia Zeylanica (Ceylon Journal of Science, Section
B—Zoology and Geology), xvii, 1-3 (1932-1933).

CoLumBuS.—Ohio State University and Ohio Academy of Science. “Ohio Journal
of Science’, xxxii, 5-6 (T.p. & c.) (1932); xxxiii, 1-4 (1933).

CoPpENHAGEN.—Det Kongelige Danske Videnskabernes Selskab. Biologiske
Meddelelser, x, 5-9 (T.p. & c.) (1932-1933); Mémoires, Section des Sciences,
Sme Serie, 111, 3-4. (fps &c:)) ((1932))2 wv, 1-2) (1983).

DusLin.—Royal Dublin Society. Economic Proceedings, ii, 29-31 (1933); Scientific
Proceedings, N.S. xx, 21-40 (1932-1933).—Royal Irish Academy. Proceedings,
Section B, xli, 9-13 (1933).

DurBAan.—Durban Museum and Art Gallery. Annual Report for Municipal Year
1931-1932 (1933).

HDINBURGH.—Royal Botanic Garden. Notes, xvi, 80 (T.p. & c.) (1932); xviii, 86
(1933); xix, 91 (1933); Transactions and Proceedings of the Botanical
Society of Edinburgh, xxxi, 1, Session 1931-32 (1932).—Royal Physical Society.
Proceedings, xxii, 3 (1933).—Royal Society of Edinburgh. Proceedings, lii,
4-5 (T.p. & c.) (1932-1933); liii, 1-3 (1933); Transactions, lvii, 2 (1933).

FRANKFURT, A. M.—Senckenbergische Naturforschende Gesellschaft. Abhandlungen,
xxxviii, 4, Teil 2 (T.p. & c.) (1933); Natur und Museum, Ixii, 9-12 (T.p. & c.)
(1932); Ixiii, 1-8 (1933).

GENEVA.—NSociété de Physique et d'Histoire Naturelle. Compte Rendu des Séances,
X1ix) 3) (CM pe Cc) 4932) laale2 933)"

Grenova.—Museo Civico di Storia Naturale. Annali, lv (1930-1932).—Societa
Entomologica Italiana. Bollettino, lxiv, 8-10 (T.p. & c.) and Supplement to
No. 9 (Memorie, xi, 1) (1932); Ixv, 1-2 and Supplement to No. 2 (Memorie,
xi, 2, Contents) (1933); 3-7 (1933).

DONATIONS AND EXCHANGES. xi

GRANVILLE.—Denison University. Journal of the Scientific Laboratories, xxvii,
pp. 47-147 (T.p. & c.) (1932); xxviii, pp. 1-154 (1933).

HaAriraAx.—Nova Scotian Institute of Science. Proceedings, xviii, 2, 1931-32 (1933).

Hatie.—Kaiserliche Leopold.-Carolin. Deutsche Akademie der Naturforsche zu
Halle. Nova Acta Leopoldina, Neue Folge, i, 2-3 (1933); Bericht, 1932-1933
(1933) F:

HampBure.—Naturwissenschaftlicher Verein. Abhandlungen, xxii, 3-4 (T.p. & c.)
(1929); xxiii, 1 (1931); Verhandlungen, Vierte Folge, iv, 1-4 (T.p. & c.)
(1930-1931).

Haritem.—NSociété Hollandaise des Sciences. Archives Néerlandaises des Sciences
exactes et naturelles, Série IIIC (Archives Néerlandaises de Physiologie de
Vhomme et des animaux), xvii, 3-4-(T.p. & c.) (1932); xviii, 1-2 (1933);
Archives Néerlandaises de Phonétique expérimentale (Organe officielle de la
Société internationale de Phonétique expérimentale), viii-ix (1933).

HEERLEN.—Geologisch Bureau voor het Nederlandsche Mijngebied. Jaarverslag,
1931 (1932).

HELsinerors.—Societas pro Fauna et Flora Fennica.—Acta Botanica Fennica, x-xi
(1931-1933); Acta Zoologica Fennica, xiii-xv (1932-1933); Memoranda, vii,
1930-1931 (1931-1932) —Societas Scientiarum Fennica. Arsbok-Vuosikirja, x,
1931-1932 (1933); Bidrag till Kannedom af Finlands Natur och Folk, |lxxxiv,
4-8 (1932-1933); Commentationes Physico-Mathematicae, vi, 16-27 (T.p. & c.)
(1932-1933) .—Societas Zoolog.-botanica Fennica Vanamo. Annales, xii (1932);
Annales Botanici, i-ii (1931-1932).

HrrosHima.—Hiroshima University. Journal of Science, Series B, Div. 1, ii, 1-7
(1932-1933); Div. 2, i, 10-14 (T.p. & c.) (1938).

Hopart.—Royal Society of Tasmania. Papers and Proceedings for the Year 1932
(1933).

HonoLtuLu.—Bernice Pauahi Bishop Museum. Bulletins 95-100; Occasional Papers,
xt, 20225 1p s & Cx 1982);

Huntineron, Long Island, N.Y.—Vanderbilt Marine Museum. Bulletin, i, 3 (1932);
iv (1933).
INDIANAPOLIS.—Indiana Academy of Science. Proceedings, xli, 1931 (1932).

Iowa City.—University of Iowa. University of Iowa Studies in Natural History,
MLV; (Os XV, 1-40 (983):

IrkutsK.—East-Siberian Geological Prospecting Trust (formerly Geological Pros-
pecting Service of the East-Siberian Region of U.S.S.R.). Records of Geology
and Mineral Resources of East-Siberia, No. 5 (1932); Transactions, Fase. 1
(1932); 1 volume in Russian (1932).

IrHAca.—Cornell University. 18 Abstracts of Theses; 7 Theses; 32 Theses (Nos.
1080-1084, 1086, 1088, 1089, 1092-1094, 1096, 1098-1104, 1106, 1107, 1109-1112,
1114, 1117, 1118, 1120, 1124-1126) (1930-1933); The George Fisher Baker Non-
Resident Lectureship in Chemistry. Vol. x (19382).

JAMAICA PLain.—Arnold Arboretum. Journal, xiii, 4 (T.p. & c.) (1932); xiv, 1-3
(1933).

JOHANNESBURG.—South African Association for the Advancement of Science.
South African Journal of Science, xxix (1932).

xii DONATIONS AND EXCHANGES.

KurAGHIKI.—Ohara Institute for Agricultural Research. Berichte, v, 3-4 (1932-
1933).

La Jorxta.—Scripps Institution of Oceanography of the University of California.
Bulletin, Technical Series, iii, 6-8 (1932); Two reprints: “Filtration Technic”
and “Mechanical Spinner for Hsmarch Cultures’ by Haldane Gee (from
Journ. of Bact., xxiv, i, July, 1932).

La PLrata.—Museo de La Plata. Obras Completas y Correspondencia Cientifica de
Florentino Ameghino, ix (Atlas) (1932).

LEWwEN.—Rijks Herbarium. Mededeelingen, No. 58A (1932).

LENINGRAD.—Académie des Sciences de VU.R.S.S. Annuaire du Musée Zoologique,
Xxxli, Beilage (1932); Bulletin, Série vii, 1931, 10 (T.p. & c.) (19381); 1932,
6-10 (1932); 19338, 1-5 (1933); Travaux du Musée Botanique, xxiv-xxv (1932) .—
Geological and Prospecting Service, U.S.S.R. Bulletin of the Information
Service of the Association for the Study of the European Quaternary at the
Geological and Prospecting Service of the U.S.S.R., Nos. 1-2 (1931-1932) ;
Problems of Soviet Geology, i, 1933, 1-3; ii, 5-6 (1933).—Lenin Academy of
Agricultural Sciences in U.S.S.R.; Institute of Plant Industry. Bulletin of
Applied Botany, of Genetics and Plant-Breeding, Series A (Plant Industry in
U.S.S.R.) (Periodical Edition), Nos. 1-2 (1932); Series i, No. 1 (1933);
Series ii, No. 1, 3 (1932); Series iii, Nos. 1 (1933), 2 (1932); Bibliographical
Contributions, No. 1 (1932); ‘Essential Oil Plants, their Cultivation and
Essential Oils’, by Prof. H. V. Wulff. Vol. i, General Part (1933).—wSociété
Russe de Minéralogie. Mémoires, 2me Série, lxi, 1-2 (T.p. & c.) (1932); lLxii,

Ae (4933)-

LikeE.—NSociété Royale des Sciences de Liége. Bulletin, i, 1932, 7-12 (T.p. & ec.)
(1932); ii, 19338, 1-5 (1933); Mémoires, 3me Série, xvii (1932).

Lincotn.—University of Nebraska, College of Agriculture, Agricultural Haperi-
ment Station. Research Bulletin 66 (1933).

LiverPooLt.—Liverpool School of Tropical Medicine. Annals of Tropical Medicine
and Parasitology, xxvi, 3-4 (T.p. & c.) (1932); xxvii, 1-2 (1933).

Lonpon.—British Museum (Natural History). A Monograph of the Recent
Cephalopoda. Pt. 2. The Octopoda (excluding the Octopodinae). By G. C.
Robson (1932); Catalogue of the Rock Collections in the Mineral Department
of the British Museum (Natural History). Part 2. America. By W. C.
Smith (1932); Hconomic Series No. 14 (1932); Index Animalium, Sectio
secunda, Pts. xxvii-xxxi (1931-1932); Insects of Samoa and other Samoan
Terrestrial Arthropoda, Pt. vi, Fasc. 7 (1931); vii, 4 (1932).—Hntomological
Society of London. Proceedings, vii, 2-3 (T.p. & ce.) (19382-1933); viii, 1
(1933); Transactions, Ixxx, 2 (T.p. & c.) (1932); Ilxxxi, 1 (1933).—Geological
Society. Geological Literature added to the Library during the Year ended
December 31st, 1931 (1932); Quarterly Journal, Ixxxviii, 4 (1st and 2nd
Halves) (T.p. & ¢.) (1932); Ixxxix, 1-3 (1933).—Linnean Society. Journal,
Botany, xlix, 328-329 (1933); Zoology, xxxvili, 258-259 (1932-1933); Pro-
ceedings, 144th Session, 1931-32, pt. 6 (T.p. & c.) (1932); 145th Session,
1932-33, pts. 1-3 (1933); Transactions, 2nd Series, Zoology, xix, 3 (1933).—
Ministry of Agriculture and Fisheries. Journal, xxxix, 8-12 (T.p. & ¢.) (1932-
1933); xl, 1-7 (1933); Report on the Work of the Land Division of the
Ministry for the Year 1932 (1933).—Royal Botanic Gardens, Kew. Bulletin

e

DONATIONS AND EXCHANGES. xliil

of Miscellaneous Information, 1932 (1933); Hooker’s Icones Plantarum, Fifth
Series, ii, 4 (T.p. & ¢.) (1933).—Royal Microscopical Society. Journal,
Series iii, lii, 4 (T.p. & ¢.) (1932); liii, 1-2 (1933) —Royal Society. Philo-
sophical Transactions, Series B, cexxi, B 480-482 (T.p. & ¢.) (1932); ccxxii,
B 483-491 (1932-1933); Proceedings, Series B, cxii, B774-779 (T.p. & Cc.)
(1932-1933); exiii; B 780-785 (T.p. & c.) (1933).—Zoological Society. List of
Fellows, April, 1932 (1932); April, 1933 (1933); Proceedings, 1932, 4 (T.p. & ¢.
for pp. 595-1084) (1933); 19383, 1-3 (T.p. & ¢. for pp. 1-541) (1933).

Los Banos, Laguna.—University of the Philippines: College of Agriculture. “The
Philippine Agriculturist”’, xxi, 7-10 (T.p. & c.) (1932-1933); xxii, 1-5 (1933).

Lunp._Kungliga Karolinska Universitetet. Lunds Universitets Arsskrift (Acta
Universitatis Lundensis), Ny Foljd, Avd. 2, xxviii, 1-2 (T.p. & ¢c.) (1932).

Lyon.—Société Linnéenne de Lyon. Bulletin Mensuel, 1re Année, Nos. 1-10
(Index) (1932); Annales de la Société Botanique de Lyon, xxxi, 3-4 (1906) ;
xxxii, 1-2 (1907); xxxiii (complete) (1908); xxxiv, 1-2 (1909); xlli, 1921
(complete) (1922).

MApison.—Wisconsin Academy of Sciences, Arts and Letters. Transactions, xxvii-
XXVili (1932-1933).

Mapras.—Madras Fisheries Department. Administration Report for the Year
1931-19382 (19383).

Mapriv.—Junta para Ampliacion de Estudios e Investigaciones Cientificas. Memoria
Correspondiente a los Cursos, 1928-9 y 1929-30 (1930); Trabajos del Museo
Nacional de Ciencias Naturales, Serie Zoologica, No. 57 (1932).—Sociedad
Espanola de Historia Natural. Boletin, xxxii, 7-10 (T.p. & c.) (1932); xxxiii,
1-3 (1933); Revista Espanola de Biologia, i, 2-3 (1932).

MANCHESTER.—Oonchological Society of Great Britain and Ireland. “Journal of
Conchology”’, xix, 8-10 (1932-1933).—Manchester Literary and Philosophical
Society. Memoirs and Proceedings, Ixxvi, 1931-1932 (1932).—Manchester
Museum. Museum Publications 102-103 (1932-1933).

MANHATTAN.—American Microscopical Society. Transactions, li, 4 (T.p. & c.)
(Index) (1932); lii, 1-3 (1933).

MAniLA.—Bureau of Science of the Government of the Philippine Islands. Thirtieth
Annual Report for Year Ending December 31, 1931 (1932); “Philippine Journal
on Sciences xlix, 3-4((ip. & cs) si932)); 1, 1-4- (ip: & c)) (1933)5 iy aes
((aL838).3

MARSEILLE.—Musée d'Histoire Naturelle de Marseille. Annales, xxiv (1931).

MELBOURNE.—‘‘Australasian Journal of Pharmacy”, N.S. xiii, 155-156 (Index)
(1932); xiv, 157-166 (1933) (From the Publisher).—Council for Scientific and
Industrial Research. Bulletin, Nos. 68, 70-76 (1932-1933); Journal, v, 4
(T.p. & c.) (19382); vi, 1-3 (1933); Pamphlets, Nos. 34-36 (1932); 37 (1933)
(Note: No 87 is also N.S.W. Department of Agriculture, Science Bulletin,
No. 40); 38-45 (1933).—Department of Agriculture of Victoria. Journal, xxx,
12 (1932); xxxi, 1-11 (1933).—Field Naturalists’ Club of Victoria. “The
Victorian Naturalist’, xlix, 8-12 (T.p.-& c¢.) (1932-1933); 1, 1-7 (1933).—
National Museum. “The Book of the Public Library, Museums and National
Gallery of Victoria, 1906-1931”, by E. la T. Armstrong and R. D. Boys (1932).—

xliv DONATIONS AND EXCHANGES.

Public Library, Museums and National Gallery of Victoria. Report of the
Trustees for 1932 (1933) —Royal Society of Victoria. Proceedings, N.S. xlv,
1-2 (T.p. & c.) (1933).—University of Melbourne. Calendar, 1933 (1932).

Mexico.—Instituto Geologico de Mexico. Anales, v (with separate Plates) (1930}.
MILWAUKEE.—Public Museum. Bulletin, ii, 4; vii, 1; xii, 2; xiii; xiv, 1 (1933).

Monaco.—Institut Océanographique de Monaco. Bulletin, No. 26 (1905); Nos.
608-610 (T.p. & ec. for Nos. 590-610) (1932); 611-633 (1933).

Montevipr0.—Museo de Historia Natural. Anales, Serie ii, iii, 3 (T.p. & ce.) (1932).

MontTrEAL.—Laboratoire de Botanique de VUniversité de Montréal. Contributions,
Nos. 22-23 (1932).

Moscow.—Limnologische Station zu Kossino Hydrometeorologischen Komitees der
U.S.S.R. Arbeiten, 15 (1932).

MuncHEenN.—Bayerische Akademie der Wissenschaften. Abhandlungen, Neue Folge,
11-16 (1932-1938); Suppl.-Band, 15 Abh. and T.p. & ec. (1932); Festrede—
“Neue Wege der beschreibenden Anatomie” von Dr. S. Mollier (1933);
Sitzungsberichte, 1932, 2-3 (T.p. & c.) (1932-1933); 1933, 1 (1933).

Nanxkine.—Metropolitan Museum of Natural History, Academia Sinica. Sinensia.
Contributions from the Metropolitan Museum of Natural History, ii, 10-12
(T.p. & c.) (19382); iii, 1-10 (1932-1933).—National Research Institute of
Geology, Academia Sinica (formerly at Shanghai). Contributions, No. 4
(1933); Memoirs, No. 12 (1931).—Science Society of China. Contributions
from the Biological Laboratory, Botanical Series, vi, 9-10 (T.p. & ce.) (1931);
vii, 1-10 (T.p. & c.) (1932); Zoological Series, viii, 1-4 (1931-1932).

NANTES.—Société des Sciences Naturelles de VlOuest de la France. Bulletin, 5me
Série, i, 1931, 1-4 (T.p. & c.) (1932).

Napies.—Stazione Zoologica di Napoli. Pubblicazioni, xii, 3 (T.p. & c.) (1933).

New Haven.—Connecticut Academy of Arts and Sciences. Transactions, xxxi,
pp. 299-425 (T.p. & c.) (1933); xxxii, pp. 1-19 (1933).—Yale University:
Peabody Museum of Natural History. Bulletin of the Bingham Oceanographic
Collection, iii, 5 (1932); iv, 3-4 (1933).

New York.—American Geographical Society. “Geographical Review”, T.p. & ec.
for xxii (1932); xxiii, 1-4 (1933).—American Museum of Natural History.
Bulletin, lili (1927); Ixiii, 4-6 (1932); Ixv (1932); Ixvii, 1 (1933); “Natural
History’, xxxii, 6 (T.p. & c.) (1932); xxxiii, 1-5 (1933).—New York Academy
of Sciences. Annals, T.p. & c. for xxxiii (1931-1932); xxxiv, pp. 1-272 (1932-
1933).

NISHIGAHARA, Tokyo.—Imperial Agricultural Experiment Station in Japan.
Journal, ii, 1-2 (1932-1933).

Osto.—Det Norske Videnskaps-Akademi i Oslo. Arbok, 1931; 1932 (1932, 1933);
Avhandlinger, I. Mat.-Naturv. Klasse, 1931; 1932 (1932, 1933); Hvalradets
Skrifter (Scientific Results of Marine Biological Research), Nos. 4-7 (1932-
1933); Skrifter, I. Mat.-Naturv. Klasse, 1931; 1932 (2 vols.) (1932, 1933).—
Université Royale. “Genetics and Cytology of the Tetraploid Form of Primula
sinensis”, by A. S. Some (From Journ. of Genetics, xxiii, 3, Dec., 1930);
“Om Planteveksten i Grensetrakter mellem Hallingdal og Valdres”, by H.
Resvoll-Holmsen (From Skrifter det Norske Videns.-Akad. i Oslo, I. Mat.-
Naturv. Klasse, 1931, No. 9) (1932).

DONATIONS AND EXCHANGES. xlv

OrrawA.—Department of Mines. Report for Fiscal Year ending March 31, 1932
(1932) ; Geological Survey of Canada. Bulletin, Nos. 69, 70 (1932); Economie
Geology Series, Nos. 11, 12 (1932); Memoir 170 (1932); Summary Report,
1932, Ai and Aii, B-D (1933).—Royal Society of Canada. Transactions, Third
Series, xxvi, Sections iv-v (1932); xxvii, List of Officers, etc. (1933); “Fifty
Years Retrospect’”—Anniversary Volume, 1882-1932 (1932).

Paris.—“Journal de Conchyliologie”’, \xxvi, 4 (T.p. & c.) (1932); Ixxvii, 1-2 (1933)
(From the Publisher) .—Museum National d’Histoire Naturelle. Archives, 6me
Série, viii (1932); Bulletin, 2e Série, iv, 3-8 (T.p. & c.) (1932); v, 1-2 (1933).—
Société Entomologique de France. Annales, ci, 3-4 (T.p. & c.) (1932); cii, 2
(1933); Bulletin, xxxvii, 1932, 15-18 (1932); xxxviii, 1933, 6-14 (1933).

Pavia.—Istituto Botanico della R. Universita di Pavia. Atti, Ser. iv, iii, 1932
(1932).

Prrpinc.—F'an Memorial Institute of Biology. Annual Report, 3rd, 1931 (1932);
4th, 1932 (1933); Bulletin, T.p. & c. for i (1929-1930); ii, 1-17 (1931); iii,
1-20 (1932).—National Geological Survey of China. Geological Bulletin, Nos.
17-22 (1931-1933).—Peking Society of Natural History and Department of
Biology, Yenching University. Peking Natural History Bulletin, vii, 2-4 (T.p.
& c.) (1932-1933); viii, 1 (1933).

PEerM.—Institut des Recherches Biologiques a VUniversité de Perm.—Bulletin, vii,
4-5 (T.p. & c.) (1930); viii, 2-5 (1932-1933); Travaux, T.p. & ec. for ii (1928-
1930); iii, 1 (T.p. & ec.) (1930-1931); iv, 1-2 (1931).

PertH.—Department of Agriculture of Western Australia. Journal, 2nd Series,
ix, 4 (T.p. & c.) (19382); x, 1-3 (1933)—Geological Survey of Western
Australia. Annual Progress Report for Year 1932 (1933).—Government
Statistician, Western Australia. Pocket Year Book of Western Australia,
1933 (1933); Quarterly Statistical Abstract, No. 250 (1928); Nos. 267-270
(1932-1933).—Royal Society of Western Australia. Journal, xvii, 1930-1931
(1931).

PHILADELPHIA.—Academy of Natural Sciences of Philadelphia. Proceedings, 1xxxiv,
1932 (19383); Review of 1932 (1933)—American Philosophical Society.
Proceedings, Ixxi, 6-7 (T.p. & c.) (1932); Ixxii, 1-3 (1933); Transactions,
N.S. xxiii, 2 (1933).—Wistar Institute of Anatomy and Biology. Bulletin
No. 8 (1933); “The Journal of Experimental Zoology”, lxiii, 2 (T.p. & c.)
(93:2) ee xvas 13s (eps ce) LIS 2-11933) aly, d-3) (Lp & c>)) (933) svat
(1933).—Zoological Society of Philadelphia. 61st Annual Report for the Year
ending February 28, 1933 (1933); Report of Laboratory and Museum of
Comparative Pathology of Zoological Society in conjunction with the 61st
Annual Report of the Society (1933).

PIETERMARITZBURG.—Natal Museum. Annals, T.p. & c. for vi (1932); vii, 2 (1933).
PirrsBurG.—Carnegie Museum. Annals, xxi (bound, complete) (1931-1933).

PLyMoutH.—Marine Biological Association of the United Kingdom. Journal,
INES 2einbbl, A (Cryo, C4@3)) (Gib) So sabe al (GER R)E
Porticit.—Laboratorio di Zoologia generale e Agraria della R. Scuola Superiore
ad’ Agricoltura. Bollettino, xxvii (1932-1933).
Prac.—Deutsche Naturwissenschaftlich-medizinische Verein fur Bohmen ‘Lotos”’
in Prag. Naturwissenschaftliche Zeitschrift “Lotos”’, Ixxx (1932)—Museum
fe)

xlvi DONATIONS AND EXCHANGES.

Nationale de Prague: Section Entomologique. Bulletin (Sbornik Entomo-
logickeho Oddeleni Narodniho Musea v. Praze), x (Nos. 72-82) (1932).—
Societas Entomologica Cechosloveniae. Acta, xxviii, 1-8 (T.p. & e.) (1931);
xxix, 1-4 (T.p. & c.) (1932).

Pretor1a.—Deparitment of Agriculture. Science Bulletin, No. 115 (1932).

Pusa.—Agricultural Research Institute. “Indian Journal of Agricultural Science”,
ii, pt. 4, pp. 345-377, 385-408 (1932); ii, pt. 5, pp. 481-454, 484-498, 531-541
(1932); ii, pt. 6, pp. 543-590, 607-637, 679-703 (1932); iii, pt. 1, pp. 1-56, 89-154;
iii, pt. 2, pp. 157-300, 320-352, 360-364, 369-376 (1933); Memoirs of the Depart-
ment of Agriculture in India, Botanical Series, T.p. & c. for xix (1933);
Entomological Series, T.p. & c. for xii (1933); Scientific Reports, 1931-32
(1933).

RIicHMOND.—Hawkesbury Agricultural College. H.A.C. Journal, xxix, 11-12 (Index)
(1932); xxx, 1-10 (1933).

Rio pE JANEIRO.—Escola Superior de Agricultura e Medicina Veterinaria. Archivos,
x, 1 (1933).—Instituto Oswaldo Cruz. Memorias, xxvi, 3 (T.p. & c.) (1932);
xxvii, 1-2 (1933).

RIVERSIDE.—University of California: Graduate School of Tropical Agriculture and
Citrus Experiment Station. Papers, Nos. 242, 247, 250, 256-258, 260-261, 263-272,
274-276 (1931-1933).

Sr. Louis.—Missouri Botanical Garden. Annals, xix, 4 (T.p. & c.) (1932); xx,
1-2 (1933).

Sr. Paut.—University of Minnesota: Agricultural Experiment Station. Bulletins
287, 288, 290-292 (1932); Technical Bulletins, 82, 84, 87 (1932).

San Dirco.—San Diego Society of Natural History. Transactions, vii, 7-23 (1932-
1933).

San FRANcIScO.—California Academy of Sciences. Proceedings, Fourth Series,
xix, 11-14 (1930-1931); xx, 1 (1931).

Sao PauLo.—Museu Paulista. Revista, xvii, 2 (1932).

Sapporo.—Hokkaido Imperial University. Journal of the Faculty of Science,
Series iv (Geology and Mineralogy), i, 3-4 (T.p. & c.) (1932); Series v
(Botany), T.p. & c. for i; ii, 1-4 (T.p. & c.) (1932-1933); Series vi (Zoology),
ii, 2-3 (1932-1933).

SEATTLE.—University of Washington Oceanographic Laboratories (formerly Puget
Sound Biological Station). Publications in Oceanography, i, 1-2 (1932-1933) ;
Publications in Oceanography, Supplementary Series, Nos. 1-5 (1931-1932).

SenpAl.—Tohoku Imperial University. Science Reports, Second Series, xii, 2B
(T.p. & c.) (1933); xv, 3 (T.p. & c.) (1932); Fourth Series, vii, 4 (T.p. & c.)
(1932); viii, 1-2 (1933).

SHARON.—Cushman Laboratory for Foraminiferal Research. Contributions, viii, 4
(1932); ix, 1-2 (1933).

Sorra.—Société Botanique de Bulgarie. Bulletin, v (1932).

STOCKHOLM.—Kungliga Svenska Vetenskapsakademien. Arkiv for Botanik, xxv,
1-3 (1932-1933); Arkiv for Kemi, Mineralogi och Geologi, xi, 1-2 (1933); Arkiv
for Matematik, Astronomi och Fysik, xxiii, 2-3 (1933); Arkiv for Zoologi,

DONATIONS AND EXCHANGES. x] vii

xxiv, 2-3 and Plates for xxiv A, No. 8 (1932-1933); 4 (T.p. & ¢c.) (1933);
xxv, 1-3 (1933); Arsbok, 1931 (1931); Handlingar, Tredje Serien, xi, 2-3
(19382); xii, 1, 3-4 (1933); Skrifter i Naturskyddsarenden, Nos. 22-24
(1932-1933).—Statens Vdatskyddsanstalt (formerly Centralanstalten for
forsoksvasendet pa jordbruksomradet). Meddelande fran Centralanstalten for
forsoksvasendet pa jordbruksomradet, Nos. 420, 422, 423 (1932); Flygblad,
Nos. 1-5 (1933); Meddelande, Nos. 1-3 (1933).

SyDNEY.—Australian and New Zealand Association for the Advancement of
Science. Report of xxist Meeting, Sydney, August, 1932 (Vol. xxi) (1933).—
Australian Museum. Annual Report of the Trustees for the Year ended 30th
June, 1932 (1933); Australian Museum Magazine, T.p. & ce. for iv (1930-1932) ;
v, 1-4 (1933); Records, xviii, 9 (1933); xix, 1 (1933).—Australian National
Research Council. “Australian Science Abstracts”, xii, 1-4 (1933).—Australian
Veterinary Association. “Australian Veterinary Journal’, viii, 6 (Index)
(1932); ix, 1-5 (1933) —Botanic Gardens. ‘A Critical Revision of the Genus
Eucalyptus”, by J. H. Maiden, I.8.0., lately Govt. Botanist of N.S.W., etce.,
T.p. & c. for Vol. viii, pts. 71-75, 1923-1928 (1933)—Commonwealth of Aus-
tralia: Department of Trade and Customs. Biological Results of the Fishing
Experiments carried on by the F.1.S. “Endeavour’’, 1909-1914, Vol. vi, 1 (1933)
(Sent by the Australian Museum, Sydney).—Department of Agriculture,
N.S.W. “Agricultural Gazette of N.S.W.”, xliii, 12 (T.p. & c.) (1932); xliv,
1-11 (1933); Live Stock Diseases Report, No. 8 (1933); Science Bulletin, Nos.
35, 38, 39, 42, 43 (1930-1933); “The Trees of New South Wales’, by R. H.
Anderson (Sydney, 1932); Veterinary Research Report, No. 6, pt. 3, December,
1931 (1933) (Sent by Veterinary Research Station, Glenfield, N.S.W.).—
Department of Mines. Annual Report for Year 1932 (1933).—Drug Houses
of Australia, Ltd. ‘Australasian Pharmaceutical Notes and News’, N.S. xi,
12 (1932); xii, 1-11 (1933) —Education Department. “Education Gazette”,
xxvi, 12 (T.p. & c.) (1932); xxvii, 1-11 (1933) —Forestry Commission, N.S.W.
Report for Year ended 31st December, 1932 (1933).—IJnstitutes of Surveyors
in Australia. “The Australian Surveyor’, iv, 4-7 (1932-1933) —Naturalists’
Society of New South Wales. ‘The Australian Naturalist’, viii, 6, 8 (1931,
1932); ix, 1-2 (1933) —Public Library of New South Wales. Annual Report
of the Trustees for the Year ended 30th June, 1932 (1933)—Royal Society
of New South Wales. Journal and Proceedings, Ixvi, 2 (T.p. & ¢c.) (1933);
Ixvii, 1 (1933) —Royal Zoological Society of New South Wales. “The Aus-
tralian Zoologist’”, vii, 45 (T.p. & ec.) (1933).—Society of Australian
Genealogists. “The Australian Genealogist”, i, 1-2 (1933)—WState Fisheries,
Chief Secretary’s Department. Annual Report on the Fisheries of N.S.W. for
the Year 1932 (1933).—Sydney University Agricultural Graduates’ Associa-
tion. “Suaga’”’, iv, 3 (1932); Issue No. 12 (May, 1933); Issue No. 13 (Sept.,
1933). —Sydney University Science Society. “Seience Journal’, xii, 1 (1933).—
Technological Museum. Six Reprints from Journ. Proc. Roy. Soc. N.S.W., 1xvi
(1932-1933), by A. R. Penfold, A. R. Penfold and F. R. Morrison, A. R. Penfold
and J. L. Simonsen and M. B. Welch (3).—‘The Medical Journal of Australia’,
LIS 2 satle 22-2 GED AccCD en GLIS2 il sas 1, 1-25 LOSS) eL9 Ss ue al-220 933)
(From the Editor).—University of Sydney. Calendar for the Year 1932
(1932); Journal of the Cancer Research Committee, iv, 4 (T.p. & c.) (1933);
v, 1-3 (1933) —Wild Life Preservation Society of Australia. Twenty-third
Annual Report, 1931-December 31, 1932 (1933).

xiviii DONATIONS AND EXCHANGES.

Tokyo.—Imperial University of Japan. Journal of the Faculty of Science, Section
iv, T.p. & c. for ii (1928-1931); iii, 2 (1933).—National Research Council of
Japan. Japanese Journal of Astronomy and Geophysics, x, 2-3 (1933);
Japanese Journal of Botany, vi, 3 (1933); Japanese Journal of Geology and
Geography, x, 1-4 (T.p. & c.) (1932-1933); xi, 1-2 (1933); Japanese Journal
of Zoology, iv, 3-5 (T.p. & c.) (1932-1933); v, 1 (1933); Report, No. 10, and
T.p. & c. for Nos. 1-10 (Vol. i) (1933); ii, 1, Apr., 1931—Mar., 1932 (1933).—
Zoological Society of Japan. Annotationes Zoologicae Japonenses, xiii, 5
(1932); xiv, 1 (1933).

Trine.—Zoological Museum. Novitates Zoologicae, xxxvii, 3 (T.p. & c.) (1932);
Xxxviii, 1-2 (1932-1933).

TRONDHJEM.—Det Konelige Norske Videnskabers Selskabs. Forhandlinger, iv,
1931 (1932); v, 1932 (1933); Skrifter, 1931 (1932); 1932 (1933); Museet:
Arsberetning, 1930 (1931); 1931 (1932); Olidsaksamlingens Tilvekst, 1930
(1931); 1931 (1932).

Tunis.—Institut Pasteur de Tunis. Archives, xxi, 2-4 (T.p. & c.) (1932-1933);
Xxii, 1-2 (1933).

UpsaLA.—University of Upsala. Zoologiska Bidrag fran Uppsala, xiv (1933).

UtrecutT.—Botanisch Museum en Herbarium van de Rijksuniversiteit. Mededee-
lingen, No. 9 (1933).

VIENNA.—Naturhistorische Museum in Wien. Annalen, xlvi (1932-1933) .—
Zoologisch-botanische Gesellschaft in Wien. Verhandlungen, Ixxxii, 1932, 1-4
(T.p. & c.) (1932); Ixxwxiii, 1933, 1-2 (1933).

WarsAw.—Panstwowe Muzeum Zoologiczne (Polish Museum of Zoology). Acta
Ornithologica, i, 1-3 (1933); Annales Musei Zoologici Polonici, ix, 20-24
(T.p. & c.) (1932-1933); x, 1-3 (1933); Fragmenta Faunistica, i, 16 (T.p. & c.)
(1932); ii, 1-6 (1933).—wSocietas Botanica Poloniae. Acta, ix, 1-4 and Supple-
ment (19382).

WASHINGTON.—American Chemical Society. Industrial and Engineering
Chemistry, xxiv, 11-12 (T.p. & c.) (1932); xxv, 1-10 (1933); Analytical
Edition, v, 1-5 (1933); News Edition, x, 20-24 (T.p. & c.) (1932); xi, 1-19
(1933).—Bureau of American Ethnology. Annual Report, 47th, 1929-1930
(1932); 49th, 1931-1932 (1933); Bulletins, 99, 106, 108-111 (1932).—Carnegie
Institution of Washington. Publications, Nos. 12, 32, 42, 46, 47, 51, 55, 57,
58, 67, 68, 75, 77, 87, 94, 106, 107, 142, 145, 146, 162, 177, 181, 197, 207, 209, 218,
231, 238, 3034, 312, 323, 324, 338, 369, 377, 410, 418, 420, 422, 425-427, 429, 434
(1904-1932); Year Book, No. 31, 1931-32 (1932).—National Academy of
Sciences. Proceedings, xviii, 11-12 (T.p. & c.) (1932); xix, 1-9 (1933).—
National Research Council. Report for the Year, July 1, 1931—June 30, 1932
(1933).—Smithsonian Institution. Annual Report of the Board of Regents
for the Year ending June 30, 1931 (1932).—U.S. Department of Agriculture.
Yearbook, 1932; 1933 (1932, 1933).—U.S. Geological Survey. Fifty-third Annual
Report of the Director for Fiscal Year ended June 30, 1932 (1932); Bulletins,
830B (T.p. & c.), 831B (T.p. & c.), 835, 836D-E (T.p. & c.), 837, 839-841, 844A-B,
849A (1932-1933); Professional Papers, 161, 166, 167, 170E (T.p. & c.), 171,
173, 174, 175A-C (1932-1933); Water Supply Papers, 640, 659A-C (T.p. & c.),
695, 697, 698, 700, 709-725 (1932-1933).—U.S. National Museum. Bulletins, 100,

DONATIONS AND EXCHANGES. xlix

Vol. vi, pts. 7-8; 100, Vol. xii (1932-1933); 158, 164 (1932); Proceedings, Ixxxi,
Arts. 4, 12-15, 17-18 (T.p. & c.) (1932-1933); Ixxxii, 1-23 (1932-1933); Report
for the Year ended June 30, 1932 (1932).

WELLINGTON.—Department of Scientific and Industrial Research: Geological Survey
Branch. Twenty-sixth Annual Report, N.S. 1931-1932 (1932); Bulletin, No.
43 (1933).—Dominion Museum. “New Zealand Journal of Science and Tech-
nology”, xiv, 3-6 (T.p. & c.) (1932-1933); xv, 1-2 (1933).—New Zealand
Institute. Transactions and Proceedings, Ixiii, 2-3 (1933).

Woops Hore.—Marine Biological Laboratory. Biological Bulletin, I1xiii, 3
GED aeGs) LOS 2) mlexdvegel ge @Lspen Gy Cah (LOSS )iew lxvend-2e G93)

Private Donors (and Authors, unless otherwise stated).

CABALLERO, EpuARDO, Mexico.—Six reprints by E. Caballero and two reprints by
E. Caballero and I. Ochoterena on Leeches (From Anales Inst. Biol. Mexico,
1-iii, 1931-1932).

CHISHOLM, EK. C., M.B., Ch.M., Comboyne, N.S.W.—‘“‘Snake Bite Anomalies” (From
“Australasian Pharmaceutical Notes and News’, May and June Issues, 1933).

CuzEens, Mrs. H., Sydney (donor).—An album of New Zealand Ferns and an
album of photos of New Zealand.

GRONBLAD, RoxF, Finland.—‘A Contribution to the Knowledge of Sub-aerial
Desmids” (From Soc. Sci. Fenn. Comment. Biol. iv, 4, 1933).

JENSEN, H. L., Sydney (donor).—“Archiv fiir Mikrobiologie’, ii, 1, pp. 136-154
(1931); Meddelande frén Centralanstalten for Forsoksvasendet pa Jordbruk-
somrddet, Stockholm, Nos. 269, 320 (1924-1927).

Meyrick, E., B.A., F.R.S., Marlborough, Wilts., England.—‘Exotic Microlepidop-
tera’, iv, 10-13 (1932-1933).

O’Dwyer, Dr. Marcaret H., Princes Risborough, Bucks., England.—‘‘Recent
Advances in the Study of the Chemistry of the Hemicelluloses of Wood.
Part i. General Review of the Chemistry of Hemicelluloses” (From Journ.
Soc. Chem. Ind., Nov. 25, 1932, Vol. li, No. 48, ‘Chem. & Ind.”, pp. 968-971).

REYCHLER, LucrIEN, Saint-Nicolas (Waes), Belgium.— ‘Bd. 31 S. 191 der
‘Mitteilungen zur Geschichte der Medizin und der Naturwissenschaften’. My
Answer. i. S.O.S. followed by ii, etc.” (Antwerp, 1933).

RuMMLER, Hans, Berlin, Germany.—“iiber die Schwimmratten (Hydromyinae)”.
(From Augustheft 1932 der Zeitschrift “Das Aquarium’) (1932).

WHITLEY, GILBERT P., Sydney.—‘Some Early Naturalists and Collectors in Aus-
tralia” (Read before the Royal Australian Historical Society, July 25, 1933).

1927
1929
1905
1906
1922
1899
1932
1927
1912

1913

1888
1925

1919

1907

1920
1929
1923
1912
1927
1923
1921
1924
1911
1932
1931
1920

1921
1926

1901
1927
1930
1905
1903
1899
1924
1932

1901
1930

1931
19338
1908

LIST OF MEMBERS, 1933.

ORDINARY MEMBERS.

*Albert, Michel Francois, ‘““Boomerang’’, Elizabeth Bay, Sydney.

Allan, Miss Catherine Mabel Joyce, Australian Museum, College Street, Sydney.
Allen, Edmund, c/o Mulgrave Mill, Gordonvale, Queensland.

Anderson, Charles, M.A., D.Sc., Australian Museum, College Street, Sydney.
Anderson, Robert Henry, B.Se.Agr., Botanic Gardens, Sydney.

Andrews, Ernest Clayton, B.A., F.G.S., 32 Benelong Crescent, Bellevue Hill.
Andrews, John, B.A., Department of Geography, Sydney University.
Armstrong, Jack Walter Trench, “Callubri’, Nyngan, N.S.W.

Aurousseau, Marcel, B.Sc.

Badham, Charles, M.B., Ch.M., B.Sc., Bureau of Microbiology, 93 Macquarie Street,
Sydney.

Baker, Richard Thomas, The Crescent, Cheltenham.

Barnard, Colin, M.Sc., Council for Scientific and Industrial Research, Division of
Plant Industry, Box 109, Canberra, F.C.T.

Barnett, Marcus Stanley, c/o Colonial Sugar Refining Co., Ltd., O’Connell Street,
Sydney.

Benson, Professor William Noel, B.A., D.Sc., F.G.S., University of Otago, Dunedin,
N.Z.

Blakely, William Faris, Botanic Gardens, Sydney.

Boardman, William, Australian Museum, College Street, Sydney.

Bone, Walter Henry, 6 Deans Place, Sydney.

Breakwell, Ernest, B.A., B.Sc., Department of Education, Box 33A, G.P.O., Sydney.

Bredero, William Adrien Lewis, Box 127, Post Office, Orange, N.S.W.

Brough, Patrick, M.A., D.Se., B.Se.Agr., Botany School, Sydney University.

Brown, Horace William, 871 Hay Street, Perth, W.A.

Brown, Miss Ida Alison, D.Sc., “Caversham”, 166 Brook Street, Coogee.

Browne, William Rowan, D.Sc., Geology Department, The University, Sydney.

Bryce, Ernest John, 47 Nelson Road, Lindfield.

Burges, Norman Alan, M.Sce., 35 Wetherell Street, Croydon.

Burkitt, Professor Arthur Neville St. George Handcock, M.B., B.Se., Medical School,
The University, Sydney.

Burns, Alexander Noble, ““Meringa’’, Fuchsia Street, Blackburn, Victoria.

Buzacott, James Hardie, Meringa (private bag), via Cairns, North Queensland

Campbell, John Honeyford, I.8.0., M.B.H., Royal Canadian Mint, Ottawa, Canada.

Campbell, Thomas Graham, ‘“Burrandong’’, 101 Lauderdale Avenue, Manly.

Carey, Miss Gladys, B.Sc., 32 Rawson Street, Epping.

Carne, Walter Mervyn, University of Tasmania, Hobart, Tasmania.

Carter, Herbert James, B.A., F.E.S., “Garrawillah’’, Kintore Street, Wahroonga.

Cheel, Edwin, Botanic Gardens, Sydney.

Chisholm, Edwin Claud, M.B., Ch.M., Comboyne, N.S.W.

Churchward, John Gordon, B.Se.Agr., Graduate College of Agriculture, The
University, Minnesota, U.S.A.

Cleland, Professor John Burton, M.D., Ch.M., The University, Adelaide, S.A.

Cochran, William Manning Patrick, B.A., c/o Bank of New South Wales, Salamoa,
New Guinea.

Colefax, Allen N., B.Se., Department of Zoology, Sydney University.

Coleman, Mrs. Edith, ‘‘Walsham’’, Blackburn Road, Blackburn, Victoria.

Cotton, Professor Leo Arthur, M.A., D.Sc., Geology Department, The University,
Sydney.

* Life Member.

1928
1900
1925

1929

1885

1930

1932
1929
1925
1928

1927
1921
1926
1920

1931
1932
1930
1914

1908
1930
1911
1886
1930

1932
1912
1911

1910
1901

19338

1925
1919
1897
1885
1928
1922

1917
1932
1911
1930
1930
1932

1907
1892

LIST OF MEMBERS. li

Craft, Frank Alfred, B.Sec., 11 Mulgray Avenue, Maroubra.

Crago, William Henry, M.D., 135 Macquarie Street, Sydney.

Cunningham, Gordon Herriot, Ph.D., Department of Agriculture, Fields Division,
Plant Research Station, P.O. Box 442, Palmerston North, N.Z.

Dakin, Professor William John, D.Sc., Department of Zoology, The University,
Sydney.

David, Sir Tannatt William Edgeworth, K.B.E., C.M.G., D.S.O., M.A., D.Sc., F.R.S.,
Burdett Street, Hornsby.

Davies, Professor Harold Whitridge, M.B., B.S., Department of Physiology, Sydney
University.

Davis, Harrold Fosbery Consett, St. Paul’s College, Newtown.

Deane, Cedric, A.M.I.E.Aust., “‘Cloyne’’, 6 State Street, Malvern, S.E.4, Victoria.

de Beuzeville, Wilfred Alexander Watt, J.P., ‘““Melamere,’’ Welham Street, Beecroft.

Dickson, Bertram Thomas, B.A., Ph.D,, Council for Scientific and Industrial
Research, Division of Plant Industry, Box 109, Canberra, F.C.T.

*Dixson, William, ‘‘Merridong’”’, Gordon Road, Killara.

Dodd, Alan Parkhurst, Prickly Pear Laboratory, Sherwood, Brisbane, Q.

Dumigan, Edward Jarrett, State School, Manly, Queensland.

Dwyer, Rt. Rev. Joseph Wilfrid, Bishop of Wagga, Wagga Wagga, N.S.W.

Edmonds, Miss Enid Mary, M.Sc., Department of Zoology, Sydney University.
*HEllis, Ralph, 2420 Ridge Road, Berkeley, California, U.S.A.

English, Miss Kathleen Mary Isabel, B.Sc., March Street, Yass, N.S.W.
Enright, Walter John, B.A., West Maitland, N.S.W.

Flynn, Professor Theodore Thomson, D.Se., Queen’s University, Belfast, Ireland.

Fraser, Miss Lilian Ross, M.Sc., ‘‘Hopetoun’’, Bellamy Street, Pennant Hills.

Froggatt, John Lewis, B.Sec., Department of Agriculture, Rabaul, New Guinea.

Froggatt, Walter Wilson, F.L.S., Young Street, Croydon.

Fuller, Miss Mary Ellen, B.Se., Council for Scientific and Industrial Research,
Box 109, Canberra, F.C.T.

Gay, Francis Joseph, B.Se., Glebe House, Reid, Canberra, F.C.T.

Goldfinch, Gilbert M., c/o Mrs. T. Savage, Archbold Road, Roseville.

Greenwood, William Frederick Neville, F.L.S., F.E.S., c/o Colonial Sugar Refining
Co., Ltd., Lautoka, Fiji.

Griffiths, Edward, B.Sc., Department of Agriculture, Raphael Street, Sydney.

Gurney, William Butler, B.Sc., F.E.S., Department of Agriculture, Raphael Street,
Sydney.

Guthrie, Tom Harrison, Whale Beach, N.S.W.

Hale, Herbert Matthew, South Australian Museum, Adelaide, S.A.

Hall, Leslie Lionel, ‘‘Haldor’, Drumalbyn Road, Bellevue Hill.

Halligan, Gerald H., F.G.S., ‘““Alameda’’, Challis Avenue, Turramurra.

Hamilton, Alexander Greenlaw, ‘‘Tanandra’’, Hercules Street, Chatswood.

Hamilton, Edgar Alexander, 16 Hercules Street, Chatswood.

Hardwick, Frederick George, B.D.S., D.D.Sc., ‘‘Wyoming’’, 175 Macquarie Street,
Sydney.

Hardy, G. H. Hurlstone, The University, Brisbane, Q.

Harris, Miss Thistle Yolette, B.Se., 93 Beach Road, Edgecliff.

Haviland, The Venerable Archdeacon F. E., Moore Street, Austinmer, South Coast,
N.S.W.

Heydon, George Aloysius Makinson, M.B., Ch.M., School of Public Health and
Tropical Medicine, The University, Sydney.

Holmes, Professor James Macdonald, B.Se., F.R.G.S., Department of Geography,
The University, Sydney.

Hossfeld, Paul Samuel, M.Se., Home Affairs Department, Canberra, F.C.T.

Hull, Arthur Francis Basset, Box 704, G.P.O., Sydney.

Hynes, Miss Sarah, B.A., “Isis”, Soudan Street, Randwick.

* Life Member.

lii

1912

1917
1932
1930
1907

1930
1933
1930

1932
1923
1924

1932
1923

1893

1922
1932

1931
1929
1932
1905
1933
1919
1932
1907
1917
1927
1919
1925
1932
1925
1930
1926

1920

1925

1913

1922

1920

1912

1920

LIST OF MEMBERS.

Irby, Llewellyn George, Forestry Department, Hobart, Tasmania.

Jacobs, Ernest Godfried, ‘‘Cambria’’, 106 Bland Street, Ashfield.

Jarvis, Edmund, Meringa (Private bag), Cairns, N. Queensland.

Jensen, Hans Laurits, McMaster Laboratory, Sydney University.

Johnston, Professor Thomas Harvey, M.A., D.Se., F.L.S., The University, Adelaide,
S.A.

Joplin, Miss Germaine Anne, B.Sc., ““Huyton’’, Wentworth Street, Eastwood.

Judge, Leslie Arthur, 36 Romsey Street, Hornsby.

Julius, Sir: George Alfred, B.Sc., B.E., M.I.Mech.E., M.I.E.Aust., 67 Castlereagh
Street, Sydney.

Kelly, Francis de Vere, 264 Elizabeth Street, Sydney.
Kendall, Mrs. W. M., M.Se. (née Williams), 5 Queen Victoria Street, Drummoyne.
Kinghorn, James Roy, Australian Museum, College Street, Sydney.

Lawson, Albert Augustus, 9 Wilmot Street, Sydney.

Lindergren, Gustaf Mauritz, Secretary, Swedish Chamber of Commerce, Pacific
House, 249 George Street, Sydney.

Lucas, Arthur Henry Shakespeare, M.A., B.Sc., ‘“Girrahween’’, William Street,
Roseville.

Mackerras, Ian Murray, M.B., Ch.M., B.Sc., Box 109, Canberra, F.C.T.

Magee, Charles Joseph, B.Sc.Agr. (Syd.), M.Sc. (Wis.), Department of Agriculture,
Raphael Street, Sydney.

*Mair, Herbert Knowles Charles, B.Se., c/o Council for Scientific and Industrial
Research, Box 109, Canberra, F.C.T.

Mann, John, Commonwealth Prickly Pear Board, Field Station, Box 49, Post
Office, Chinchilla, Queensland.

Martin, Donald, B.Se. c/o University of Tasmania, Hobart, Tasmania.

Mawson, Sir Douglas, D.Sc., B.E., F.R.S., The University, Adelaide, S.A.

Maze, Wilson Harold, 39 Lucas Road, Burwood.

McCarthy, Timothy, Department of Agriculture, Raphael Street, Sydney.

McCulloch, Robert Nicholson, B.Sec.Agr. (Syd.), B.Se. (Oxon.), 36 Manning Road,
Double Bay.

McDonnough, Thomas, L.S., 15 Hamilton Street, Coogee.

McKeown, Keith Collingwood, Australian Museum, College Street, Sydney.

McKie, Rev. Ernest Norman, B.A., The Manse, Guyra, N.S.W.

McLuckie, John, M.A., D.Sc., Botany Department, The University, Sydney.

McNeill, Francis Alexander, Australian Museum, College Street, Sydney.

Messmer, Pearl Ray (Mrs. C. A.), Treatts Road, Lindfield.

Mitchell, Miss Dora Enid, B.Se., ‘“Frensham’’, Mittagong, N.S.W.

Munch-Petersen, Erik, Ph.B., M.Sc. (Haunensis), M.I.F., 31 Lytton Street, North
Sydney.

Mungomery, Reginald William, c/o Sugar Experiment Station, Bundaberg, Queens-
land.

Musgrave, Anthony, F.H.S., Australian Museum, College Street, Sydney.

Newman, Ivor Vickery, M.Se., Ph.D., F.R.M.S., F.L.S8., “Tip Tree’, Kingsland
Road, Strathfield.

Newman, Leslie John William, F.E.8., ‘“‘Walthamstowe’’, 5 Bernard Street, Clare-
mont, W.A.

Nicholson, Alexander John, D.Sc., F.E.S., Council for Scientific and Industrial
Research, Box 109, Canberra, F.C.T.

Noble, Robert Jackson, B.Sc.Agr., Ph.D., Department of Agriculture, Raphael
Street, Sydney.

North, David Sutherland, c/o Colonial Sugar Refining Co., Ltd., Broadwater Mill,
Richmond River, N.S.W.

O’Dwyer, Margaret Helena, B.Sc., Ph.D., Forest Products Research Laboratory,
Princes Risborough, Bucks., England.

* Life Member.

LIST OF MEMBERS. lili

Oke, Charles George, 56 Chaucer Street, St. Kilda, Victoria.

Oliver, Walter Reginald Brook, F.L.S., F.Z.S., Dominion Museum, Wellington, N.Z.

Osborn, Professor Theodore George Bentley, D.Sc., F.L.S., Department of Botany,
The University, Sydney.

Osborne, George Davenport, D.Sc., Geology Department, The University, Sydney.

Perkins, Frederick Athol, B.Sc.Agr., Biology Department, University of Queensland,
Brisbane, Q.

Phillips, Montagu Austin, F.L.S., F.E.S., 57 St. George’s Square, London, S.W.,
England.

Pincombe, Torrington Hawke, B.A., ‘““Mulyan’’, Beta Street, Lane Cove, Sydney.

Priestley, Professor Henry, M.D., Ch.M., B.Sc., Medical School, The University,
Sydney.

Pulleine, Robert Henry, M.B., Ch.M., 163 North Terrace, Adelaide, S.A.

Raggatt, Harold George, B.Sce., Geological Survey, Department of Mines, Sydney.

Roberts, Frederick Hugh Sherston, M.Se., Department of Agriculture and Stock,
Animal Health Station, Yeerongpilly, Brisbane, Q.

Robertson, Rutherford Ness, 15 The Boulevarde, Lewisham.

Roughley, Theodore Cleveland, B.Sc., F.R.Z.S., Technological Museum, Harris
Street, Sydney.

Rupp, Rev. Herman Montagu Rucker, B.A., The Rectory, Woy Woy, N.S.W.

Salter, Keith Eric Wellesley, B.Sc., ““Hawthorn’’, 48 Abbotsford Road, Homebush.

*Scammell, George Vance, B.Sce., 7 David Street, Clifton Gardens.

Selby, Miss Doris Adeline, M.Sc., ‘“Marley’’, Werona Avenue, Gordon.

Sherrard, Mrs. Kathleen Margaret, M.Sc., 16 Shellcove Road, Neutral Bay.

Smith, G. P. Darnell, D.Se., F.1.C., F.C.S., Botanic Gardens, Sydney. :

Smith, Jacob Harold, M.Sc., N.D.A., Department of Agriculture and _ Stock,
Atherton, N. Queensland.

Smith, Thomas Hodge, Australian Museum, College Street, Sydney.

Smith, Miss Vera Irwin, B.Se., F.L.S., 13 Upper Cliff Road, Northwood.

Stanley, George Arthur Vickers, B.Sc., ‘“‘Clelands’’, 33A Battery Street, Clovelly.

Stead, David G., ‘‘Boongarre’’, Pacific Street, Watson’s Bay.

Stokes, Edward Sutherland, M.B., Ch.M., Metropolitan Water, Sewerage and
Drainage Board, 341 Pitt Street, Sydney.

*Sulman, Miss Florence, ‘‘Burrangong’’, McMahon’s Point.

Sussmilch, C. A., F.G.S., Hast Sydney Technical School, Darlinghurst, Sydney.

Taylor, Frank Henry, School of Public Health and ‘Tropical Medicine, The
University, Sydney.

Tillyard, Robin John, M.A., D.Sc., F.R.S., F.L.S., F.E.S., C.M.Z.S., Chief Common-
wealth Entomologist, Canberra, F.C.T.

*Troughton, Ellis Le Geyt, Australian Museum, College Street, Sydney.

Turner, A. Jefferis, M.D., F.E.S., Wickham Terrace, Brisbane, Q.

Turner, Rowland E., F.Z.S., F.E.S., c/o Standard Bank of South Africa, Adderley
Street, Cape Town, South Africa.

Veitch, Robert, B.Sc., F.E.S., Department of Agriculture, Brisbane, Queensland.
Vickery, Miss Joyce Winifred, M.Sc., 6 Coventry Road, Homebush.

Walker, Commander John James, M.A., F.L.S., F.E.S., R.N., ‘“‘Aorangi’’, Lonsdale
Road, Summertown, Oxford, England.

Walkom, Arthur Bache, D.Sc., Science House, Gloucester and Essex Streets, Sydney.

Ward, Melbourne, ‘‘Pasadena’’, Cross Street, Double Bay.

Wardlaw, Henry Sloane Halcro, D.Sc., Physiology Department, The University,
Sydney.

Waterer, Arthur S., 2 Dickson Street, Haberfield.

*Waterhouse, G. Athol, D.Se., B.E., F.E.S., Science House, Gloucester and Essex
Streets, Sydney.

Waterhouse, Lionel Lawry, B.E., ‘Rarotonga’, 42 Archer Street, Chatswood.

* Life Member.

liv

1927

1911
1926
1926
1933
1932
1903
1925
1933
1910

1923

1923

1888
1902
1932
1902

LIST OF MEMBERS.

Waterhouse, Walter Lawry, D.Sc.Agr., M.C., D.I.C. (Lond.), Faculty of Agriculture,
Sydney University.

Watt, Professor Robert Dickie, M.A., B.Sc., University of Sydney.

Weekes, Miss Hazel Claire, D.Sc., 73 Ocean Avenue, Edgecliff.

*Whitley, Gilbert Percy, Australian Museum, College Street, Sydney.

Willings, Horace John, B.A., Department of Zoology, Sydney University.

Woodhill, Anthony Reeve, B.Sc.Agr., Department of Zoology, Sydney University.

Woolnough, Walter George, D.Sc., F.G.S., Canberra, F.C.T.

Wright, Fred, c/o Messrs. Elliott Bros., Ltd., O’Connell Street, Sydney.

Wright, Gilbert, Faculty of Agriculture, Sydney University.

Wymark, Frederick, 89 Castlereagh Street, Sydney.

HONORARY MEMBERS.
Hill, Professor J. P., Institute of Anatomy, University of London, University
College, Gower Street, London, W.C.1, England.
Wilson, Professor J. T., LL.D., M.B., Ch.M., F.R.S., Department of Anatomy, the
New Museums, Cambridge, England.

CORRESPONDING MEMBERS.
Bale, W. M., F.R.M.S., 63 Walpole Street, Kew, Melbourne, Victoria.
Broom, Robert, M.D., D.Sc., F.R.S., Bothaville, O.F.S., South Africa.
Dun, William S., Geological Survey, Department of Mines, Sydney.
Meyrick, Edward, B.A., F.R.S., F.Z.5., Thornhanger, Marlborough, Wilts., England.

* Life Member.

so 08 o™
Wg ac “y
Ve Ry Qe ‘a?

MLIBRARY|I-—

. > Oee, 3
‘ ™

INDEX Sera
° r y +
(1933.)
(a) GENERAL INDEX.
Address, Presidential, 1. Burkitt, Prof. A. N. St. G. H., elected

Alteration of Rules, 1.

Anderson, C., elected a Vice-President,
xxix—The Fossil Mammals of Australia,
1x

Andrews, J., and W. H. Maze, Seasonal
Incidence and Concentration of Rainfall
in Australia, 121—-Some Climatological
Aspects of Aridity in their Application
to Australia, 105.

Antarctic Continent, section placed under
authority of Commonwealth, ii.

Aridity, some Climatological Aspects of,
in their Application to Australia, 105.
Australia, Seasonal Incidence and Concentra-
tion of Rainfall in, 121—Some Climato-
logical Aspects of Aridity in their

Application to, 105.

Australian and New Zealand Association
for the Advancement of Science, 2ist
Meeting held, i.

Australian, Coleoptera. Notes and New
Species, 159 — Diptera, Miscellaneous
Notes on, 408—Diptera, Notes on, 74—
Hesperiidae, Notes and Descriptions of
New Forms, 461—Lepidoptera, Revision
of. Oecophoridae, 80—Oyster, the Life
History of the, 279—Proctotrypidae, a
New Genus and Species of, 275—Snakes
and Lizards, on the Distribution, Habitat
and Reproductive Habits of certain
European, and with Particular Regard to
their Adoption of Viviparity, 270.

Balance Sheets for Fourteen Months ending
28 February, 1933, xxvi-xxviii.

Bancroft, Thos. L., Some Further Cbserva-
tions on the Rearing of Ceraiodus, 467—
reference to death, xxxv—see Exhibits.

Brough, P., congratulations to, x—The
Life History of Grevillea robusta (Cunn.), 33.

Brown, Ida A., Linnean Macleay Fellow in
Geology, The Geology of the South Coast
of New South Wales, with Special Reference
to the Origin and Relationships of the
Igneous Rocks, 334—Summary of year’s
work, vi—Retirement from Fellowship,
Vil.

Burges, N. Alan, congratulations to, xxxi—
appointed Linnean Macleay Fellow of the
Society in Botany, 1934-35, xxxv.

President, xxv.

Burton, Florence R. W., elected a member,
XXXV.

Carter, H. J., Australian Coleoptera. Notes
and New Species, 159.

Casuarinas, the Coccidae of the, 363.
Catalogue of Scientific and Technical
Periodicals, issue of Supplement to, ii.
Ceratodus, Some Further Observations on

the Rearing of, 467.

Cheel, E., appointed to National Park Trust,
1i—elected a Vice-President, xxix—see
Exhibits.

Chisholm, E. C., Useful Coccinellidae found
on the Comboyne Plateau, 405.

Coastal Tablelands and Streams of New
South Wales, 437.

Cobb, Dr. N. A., obituary notice, iv.

Coccidae of the Casuarinas, 363.

Coccinellidae, Useful, found on the Comboyne
Plateau, 405.

Colefax, Allen, see Dakin, Prof. W. J., and
A. Colefax—see Exhibits.

Coleman, Mrs. Edith, elected a member,

DOO:

Coleoptera, Australian. Notes and New
Species, 159.

Comboyne Plateau, Usefu Coccinellidae

found on the, 405.

Corynebacteria as an Important Group of
Soil Microorganisms, 181.

Craft, Frank A., Linnean Macleay Fellow
in Geography, The Coastal Tablelands and
Streams of New South Wales, 437—The
Surface History of Monaro, N.S.W.,
229—Summary of year’s work, vii—re-
appointed, 1933-34, vui—reappointed,
WOS4=35,) Sexe

Dakin, Prof. W. J.—see Exhibits.

Dakin, Prof. W. J., and A. Colefax, The
Marine Plankton of the Coastal Waters
of New South Wales, 1. The Chief
Planktonic Forms and their Seasonal
Distribution, 186—see Exhibits.

David, Sir T. W. E., congratulations to, xxx.

Diptera, Australian, Miscellaneous Notes
on, 408—-Australian, Notes on, 74.

Dixson, T.S., obituary notice, ili.

lvi INDEX.

Dodd, Alan P., A New Genus and Species of
Australian Proctotrypidae, 275.

Donations and Exchanges, xxix, XXxi-xxXxv.

Drosera peltata and D.auriculata, Vegetative
Reproduction in, 245.

Dun, W.S., elected a Corresponding member,
is

Edmonds, Enid, congratulations to, xxx1.
Elections, xxv, XXiX, XXXI-XXXIll, XXXvV.
Exchange relations, 1.

Exhibits :

Bancroft, T. L., see under Whitley, G. P.

Colefax, A. N., see under Dakin, Prof.
W. J., and A. N. Colefax.

Dakin, Prof. W. J., two spider crabs,
male and ripe female, from Corner
Inlet, Victoria, xxx1.

Dakin, Prof. W. J., and A. N. Colefax, a
plankton catch and note thereon, xxxvi.

Fraser, Lilian, specimens of Duictydium
rutilum, XXXVIi.

Jacovs, E. G., a flower spike of the
“Common Sun Orchid”, Thelymitra
ixioides Swartz; three flowers of Cala-
denia alba R.Br., xxxiil.

Joplin, Germaine A., various plants from
the Hartley District, xxxil.

McLuckie, J., a preliminary note on
Cryptanthemis Slatert Rupp, xxxiv.

Messmer, Mrs. C. A., specimens of a new
species of Pterostylis from Fitzroy Falls,
sxeKeKIE

Roughley, T. C., specimens of yearling
Sea Trout (Salmo trutta) from River
Plenty, Tasmania, xxx—specimens of
an Isopod Crustacean (Cirolana corpu-
lenta) from sharks and rays caught off
Stockton Beach, xxx.

Rupp, Rev. H. M. R., sketches of Australian
Orchids, xxxii—specimens of Cryptan-
themis Slateri, xxxiv — Cleisostoma
Nugentii Bailey in flower from Proser-
pine, N. Queensland; enlarged photo-
graphs of Cryptanthemis Slatert Rupp ;
eoloured sketches of Australian orchids,

XXXVI.
Waterhouse, G. A., two _ butterflies,
Taractrocera ina and Padraona tanus

nihana, from Hayman Is., Queensland,
xxxii—pupae of Heteronympha cordace
reared from eggs laid by captive females,
xxxv—-coloured drawings of a pupa
found at Mt. Kosciusko and of the
variable larvae, xxxv—the early stages
of Hesperilla chaostola and FHesp.
donnysa, XXXvIl.

Whitley, G. P., a right-hand male and a
left-hand female of the Four-eyed Fish
(Anableps dower Gill) from Honduras ;
a cranium of a Catfish (Neoarius australis
Gunther) from Moreton Bay, Queens-
land, xxx—a Long-finned Eel (Anguilla
reinhardtii Steindacher) from Misima

Island, Papua, xxxiii—on behalf of
Dr. T. L. Bancroft, a series of specimens
of Ceratodus, xxxv.

Fraser, Lilian R., Linnean Macleay Fellow
in Botany, An Investigation of the Sooty
Moulds of New South Wales. Historical
and Introductory Account, 375—The
Mycetozoa of New South Wales, 431—
Summary of year’s work, viii—reappointed,
1933-34, viii—reappointed, 1934-35, xxxv—
see Exhibits.

Froggatt, W. W., The Coccidae of the
Casuarinas, 363.

Genus Pterostylis R.Br., A new Scheme of
Classification, with Notes on the Distribu-
tion of the Australian Species, 421.

Geology of the South Coast of New South
Wales, with special Reference to the
Origin and Relationships of the Igneous
Rocks, 334.

Grevillea robusta (Cunn.), the Life History of,
33.

Guthrie, T. H., elected a member, xxix.

Hamilton, H. W., reference to death, xxix.

Hardy, G. H., Miscellaneous Notes on
Austrahan Diptera, 408.

Hartley District, the Petrology of the.
The Metamorphosed Gabbros and
Associated Hybrid and Contaminated
Rocks, 125.

Hesperiidae, Australian. Notes and
Descriptions of New Forms, 461.
Holmes, Prof. J. Macdonald, elected a

member of Council, xxv.

Investigation of the Sooty Moulds of New
South Wales. Historical and Introductory
Account, 375.

Iredale, Tom, and T. C. Roughley, The
Scientific Name of the Commercial Oyster
of New South Wales, 278.

Jacobs, E. G., see Exhibits.

Jensen, H. L., Macleay Bacteriologist,
Corynebacteria as an Important Group
of Soil Microorganisms, 181—Summary
of year’s work, v.

Joplin, Germaine A., congratulations to,
xxxi—The Petrology of the Hartley
District. The Metamorphosed Gabbros

and Associated Hybrid and Contaminated
Rocks, 125—see Exhibits.
Judge, L. A., elected a member, xxxiil.

Kosciusko Region, Mount, Mayflies of the.
Intreduction and Family Siphlonuridae, 1.

Lepidoptera, Australian.
Oecophoridae, 80.
Life History of Grevillea robusta (Cunn.), 33.

Revision oi,

INDEX.

Life History of the Australian Oyster (Ostrea
commercialis), 279.
Linnean Macleay Fellowships, 1933-34, re-

appointments and appointment, vili—
1934-35, applications invited, xxxiii,
XXxiv — reappointments and appoint-

ment, XXXv.

Lizards, Certain European and Australian
Snakes and, on the Distribution, Habitat
and Reproductive Habits of, with Particular
Regard to their Adoption and Viviparity,
270.

Lord, Clive, reference to death, xxxil.

Malloch, J. R., Notes on Australian Diptera,
74.

Mammals, Fossil, of Australia, ix.

Marine Plankton of the Coastal Waters of
New South Wales. The Chief Planktonic
Forms and their Seasonal Distribution, 186.

Mayflies of the Mount Kosciusko Region.
Introduction and Family Siphlonuridae, 1.

Maze, W. H., elected a member, xxxiii—
see Andrews, J., and W. H. Maze.

McAlpine, D., obituary notice, iv.

McLuckie, J., see Exhibits.

Messmer, Mrs. Pearl R., A New Species of
Pterostylis, 429—-see Exhibits.

Miscellaneous Notes on Australian Diptera,
408.

Monaro, New South Wales,
History of, 229.

Mycetozoa of New South Wales, 431.

the Surface

Native flora, standing committee appointed
to consider what might be done to
popularize, 1.

New Genus and _ Species
Proctotrypidae, 275.

Newman, I. V., appointed Linnean Macleay

of Australian

Fellow in Botany, 1933-34, viii—re-
appointed, 1934-35, xxxv.

New Species of Pterostylis, 429.

North Queensland Naturalists’ Club,

announcement of proposed camp at Low
Island, xxxiv.

Notes on Australian Diptera, 74.

Notes on New South Wales and Queensland
Orchids, 223.

Notes on the Australian Species of the
Family Paussidae (Coleoptera), 396.

On the Distribution, Habitat and Repro-
ductive Habits of certain European and
Australian Snakes and Lizards, with
Particular Regard to their Adoption of
Viviparity, 270.

On the Production of Fertile Hybrids from
Crosses
Emmer Wheats, 99.

Orchids, New South Wales and Queensland,
Notes on, 223.

Osborn, Prof. T. G. B., elected a Vice-
President, xxix.

between Vulgare and Khapli

lvii

Oyster, Australian, the Life History of the,
279—Commercial, of New South Wales,
the Scientific Name of the, 278.

Paussidae (Coleoptera), Notes on the Aus-
tralian Species of the Family, 396.

Petrology of the Hartley District. The
Metamorphosed Gabbros and Associated
Hybrid and Contaminated Rocks, 125.

Plankton, Marine, of the Coastal Waters of
N.S.W. The Chief Planktonic Forms and
their Seasonal Distribution, 186.

Presidential Address, i.

Proctotrypidae, Australian, a New Genus
and Species of, 275.

Pterostylis, a New Species of, 429—R.Br.,
the Genus (Orchidaceae), 421.

Rainfall in Australia, Seasonal Incidence and
Concentration of, 121.
Revision of Australian

Oecophoridae, 80.

Roughley, T. C., The Life History of the
Australian Oyster (Ostrea commercialis),
279 — see Iredale, Tom, and T. C.
Roughley—see Exhibits.

Rupp, Rev. H. M. R., Notes on New South
Wales and Queensland Orchids, 223—
The Genus Pterostylis R.Br. (Orchidaceae).
A New Scheme of Classification, with
Notes on the Distribution of the Aus-
tralian Species, 421—see Exhibits.

Lepidoptera.

Sales Tax, removal of, from educational
books, ii.

Scientific Name of the Commercial Oyster
of New South Wales, 278.

Seasonal Incidence and Concentration of
Rainfall in Australia, 121.

Sloane, T. G., obituary notice, lil.

Sloane, the late T. G., Notes on the Australian
Species of the Family Paussidae (Cole-
optera), 396.

Snakes and Lizards, certain European and
Australian, on the Distribution, Habitat
and Reproductive Habits of, with Particular
Regard to their Adoption of Viviparity,
270.

Soil Microorganisms, Corynebacteria as an
Important Group of, 181.

Some Climatological Aspects of Aridity in
their Application to Australia, 105.

Some Further Observations on the Rearing
of Ceratodus, 467.

Sooty Moulds of New South Wales, an
Investigation of the. Historical and Intro-
ductory Account, 375.

South Coast of New South Wales, the
Geology of the, with special Reference to
the Origin and Relationships of the
Igneous Rocks, 334.

Surface History of Monaro, N.S.W., 229.

Sussmilch, C. A., elected a member of Council,
xxyV.

lviii

INDEX.

Tillyard, R. J., The Mayflies of the Mount

Kosciusko Region, Introduction and
Family Siphlonuridae, 1

Turner, A. J., Revision of Australian
Lepidoptera. Oecophoridae, 80.

Vegetative Reproduction in Drosera peltata
and D. auriculata, 245.

Vickery, Joyce W., congratulations to, xxxi—
Vegetative Reproduction in Drosera peltata
and D. auriculata, 245.

Wardlaw, H.S. H., elected a Vice-President,
xxix—Presidential Address of, reprinted
in Annual Report of Smithsonian Institu-
tion, ll.

Waterhouse, G. A., Australian Hesperiidae.
Notes and Descriptions of New Forms,

461—elected Hon. Treasurer, xxix—see
Exhibits.

Waterhouse, W. L., On the Production of
fertile Hybrids from Crosses between
Vulgare and Khapli Emmer Wheats, 99.

Weekes, H. Claire, Linnean Macleay Fellow
in Zoology, On the Distribution, Habitat
and Reproductive Habits of certain
European and Australian Snakes and
Lizards, with particular regard to their
Adoption of Viviparity, 270—Summary
of year’s work, vii—reappointed, 1933-34,
viii.

Wheats, On the Production of fertile Hybrids
from Crosses between Vulgare and Khapli
Emmer, 99.

Whitley, G. P., see Exhibits.

Willings, H. J., elected a member, xxxi.

Wright, Gilbert, elected a member, xxix.

(b) BIOLOGICAL INDEX.

New names are printed in SMALL CAPITALS.

Acacia sp. .. 364 Antennaria elaiophila .. 377 Arthropterus denudatus
Acanthometron sp. ao dete! ericophila i oS 397, 400
Acanthorhynchus tenul- Robinsoni -. 385 dohrni . 401
ROBO bg 0 oo oo og WS) scoriadea go oa Gee) elongatulus . 402
Acartia centrura Eteata ase mei semi-ovata OU ERO SO. foveicollis . 400
clausi . 201,207,218 Antennularia scoriadea foveipennis .. 404
danae so) og, AUG 385-7, 389, 392 geminus .. , .. 401
Acrocalanus gibber 204 Anthochaera chrysoptera 65 hopei . 397, 400
gracilis .. .. .. 205,218 Apaustus dolon .. . 463 howitti . 397, 401
Adelium flavicornis .. 174 Apiocera asilica . .. 416 humeralis . 401
FLAVITARSIS a lize fuscicollis .. 416 kingi .. 400
ruptum 174 maerens .. .. 416 latipennis -. 403
striatum . so UG: MARITIMA .. 416 LONGICOLLIS . 402
Aegiceras majus . 322 vulpes .. 416 macleayi . OU,
Aepyprymnus Xvill ee citri .. 384 mastersi .. 397, 400-1
Aithaloderma citri .. 384 salicinum . So Da) Oso OF moretoni.. 5) Soo GOI
Alecanopsis casuarinae .. 364 Apogonops.. .. .. 203, 210 nigricornis 400-1
grandis . 365 anomalus.. So. oo P4lles occidentalis - 400, 402
tenuis .. 364 Arbutus unedo 5.5 Ghd occidentalis var.
Alternaria sp. 387-8 Archarthropterus . 400 ORIENTALIS .. 402°
AMELETOIDES 5 . 4,5 Archizonurus securus XVili ominosus gu erg) ADI
LACUS-ALBINAE PON WHER 6 Arcyria cinerea . .. 434 ovicollis .. 60 By BOY)
Ameletopsis Bd) 99.0) ete denudata .. 434: parallelocerus . 399, 400
Ameletus flavitinctus oaak, 1 ferruginea .. 434 pervicax .. .. 401
ornatus oom tbl glauca oo) Gey piceus We . 397
OUTS 66° 60 oo 40 1S incarnata. -. 434 PLANICORNIS . 399, 402
subnotatus ogy aly incarnata var. ‘fulgens .. 434 rockhamptonensis .. 397, 399
Amenia eens OLA OTN Ts Ova cth C7 insignis .. 434 secedens .. . 401
chrysame. . 75, 77 nutans -. 435 sphinx . 401
dubitalis .. 76-7 Oerstedtil Oo eign Cae) suspectus.. . 401
imperialis 75,77 Arthropterus Bo 06 OW, ZOO turneri Ran) asst a ecaaont: 40)
leonina i 74, 77 ambitiosus eo au CO waterhousei .. 397, 399, 401
nigromaculata. 76-7 angulatus .. 401 westwoodi . 397, 400, 402
SEXPUNCTATA 76-7 articularis .. 400 wilsoni a0 OO
Anableps dowei .. 9 XOX bisinuatus -. 400 Asbolisia sp. 387-8
Anaxo cylindricus so Oe brevicollis 399, 403 Aspidiotus cingulatus . 363
TENUICORNIS . 176 brevis .. 397 Asterionella japonica
Androsus brevis 66 7/33 CONSTRICTICEPS ee OS 192, 194-5, 216
HACKERI .. 5 783 CRIBROSUS . 400, 403 Atichia glomerulosa
violaceus : uence maladie eylindricollis See CO 376, 381, 383, 389, 392
Anguilla reinhardtii : XXXIil eylindricus . 400 Atlanta sp.. 5 oo) (oo WY
Anodonta . 325 daemelianus . 401 Aurelia caerula .. 3 IGS}

&

AUSTROSERPHUS.. so, PACKS
ALBOFASCIATUS da P4tKs}
Bacteriastrum sp... 191, 216
Bacterium herbicola.. .. 185
Badhamia capsulifera .. 432
papaveracea .. 432
utricularis Hee Gre BP
Banksia integrifolia .. 322, 364
BARYZANCLA Rulon bien!
DYSCLYTA 81
ITHYGRAMMA .. 81
Beroe sp. ah. dig ihe)
Bibla anisomorpha . 464
Biddulphia mobiliensis
191-2, 216
regia.. 191-2
JO, do | Oe 216
Bolina chuni 198
Borkhausenia 88
ALBIPECTINATA 93
BASILEUCA 96
CENTROSTICHA 98
CHALCOTEUCTA 92
CNECOCRANA 95
cyclozona 93
FLAVIPUNCTA .. 95
GYPSODES 94
GYPSOMICTA 92
hemileuca 90
HOMOPELA 9]
IULOPHYLLA 94
LECHRIOGRAMMA 97
LECHRIOMOCHLA 93
LEPTOCNECA 96
LEPTOPHYLLA . 90
LISSOPTERA 96
minutella.. 88
MISELLA .. 94
NEPHOTYPA 98
NIGRIPUNCTA .. 93
nubifera .. 93
oxypeuces 92
PELOPHANES 97
PERIGRAPTA 92
PHANEROSTICTA 9]
PSARITIS .. 4 97
SERRULIFERA .. 94
TANYTRICHA 91
TRICHOCEROS .. 98
VERNILIS .. 95
Botrytis cinerea 392
Brachysiphon 69
Bucephalus 325
haimeanus 325
polymorphus .. : 326
Bulgaria polymorpha 383
Burramys XViil
parvus XVll
Byallius andersoni 171
angustatus 170-1
kosciuskoanus 171
laticollis .. WZ/AL
mastersi .. Nez
OBERONIUS 170-1
ovensensis 170-1
punctatus 171
reticulatus 170-1
revolutus 171

INDEX.

Caladenia alba XXXII
alpina a LO . 227
dilatata var. concinna.. 228

Calanus brevicornis

xxxvi, 203, 214
darwinii » 203
finmarchicus . 203
gracilis -- 203
minor : 203- 4
patagoniensis .. es cOS
pauper ae 218
tenuicornis 204
vulgaris 204
Caldariomyces Fumago " 380- 1
Caleana minor 5 pale
Nublingii a0 Pai
Callineda pean diiertal a. 406
Calocalanus plumulosus .. 205
Campinotus intrepidus .. 365
Candace ethiopica eS
pectinata .. 218
Candacia bipinnata . 206
ethiopica . -- 206
pachydactyla .. .. 206
aoe ae a0 oo AUG
truncata . ake a6 ZAVE
Capnodium ar araucariae 385
armeniaceae % oo Oreld
australe. wie! ac. 384-5
eallitris 385-6
casuarinae 385
citri . _ 377, 384
citricolum 385- 6, 390-2
elongatum . 384
javanicum . 390
meridionale oS).
neril oop
Persoonii : AN Fen A Oe
salicinum 377- 8, 380-2, 384-5,
387, 390, 392
stellatum : . 390
scoriadeum 55 Orel
Walteri ie . 384
Carcharhinus macrurus 5 BOX
Carcharias arenarius.. .. Xxx
Carcharodon carcharias .. xxx
Cardium sre Ne O2zo
Castiarina BOOYANIA 162
hilaris var. INFASCIATA 163
INUSITATA 160
MAGNETICA 162
RADIANS .. .. 160
SUBNOTATA a9
SUBTINCTA 159

Casuarina Cambagei
364, 368, 370

distyla . 365
Cunninghamiana 5 8 Bae
humilis See Od
lepidophloia 363-5, 368
Luehmanni ..367, 370, 372
quadrivalvis 370, 372-4
rigida é . 365
sp. . 364
stricta , 372-3
suberosa .. 364- 5, 368, 374
torulosa : 5 ne Ue

lix

CASUARINALOMA.. 368
leal : 368
Centropages bradyi - 205
calaninus ss 200
chierchiae 32 ZOD
furcatus .. - 2065
orsinil Ar, PSUs)
violaceus «2 206
Cerataulina sp. .. 216
Ceratiomyxa frutic ulosa - 432
Ceratium furca .. . 196, 218
fusca so, LLS
fusus 196
palmatum 196
platycorne 196
ranipes Me 196
tripos : 196, 218
Ceratodus xxxv, 467
Cerioides ornatus . 420
subarmatus é . 420
Ceroplastes ceriferus .. 387, 392
rubens . 387, 393
Chaetoceras 194-5
affinis 191
curvisetum 191
debilis 191
didymum 191
Lorenzianus 191
Peruvianus i /oe) OIL
sociale > LOT 216
sp. ced) ar 2G
Chaetothyrium lepernanes 385
Chalcides sepoides 5 Pat le
tridactylus . 273
Chariotheca doddi (var.) .. 171
striato-punctata 171
Chlorella pacifica . 309
Chrysopilus fascipennis .. 408
Cienkowskia reticulata 2 BR:
Cirolana corpulenta .. ExSkX
Cisseis pulchella 163
SAPPHIRA 163
westwoodi Be) LG:

Cladosporium herbarum
381-3, 387-9, 392
Clastoderma Debaryanum 433

Clausocalanus arcuicornis 205
furcatus .. 5 3 205
Cleisostoma Nugentii XXXV1
Climacodium bicodenvers 216
frauenfeldianum . 192, 216
Climacograptus . - 335
Coccinella repanda - 405
Codonellopsis ostenfeldti
198, 218
Coelophora inequalis -. 406
veranioides . 406
Coleroa Straussii Fa nate!
Collema glomerulosa - 381
Colloderma oculatum 433
Collozoum inerme 198
Coloburiscus 4, 5, 20
GIGANTEUS Denil
haleuticus 20, 22
humeralis 20 —2,
MUNIONGA 22, 29

lx

Comatricha elegans 433
irregularis 433
laxa.. 433
nigra ae 433
pulchella var. “fusca 433
typhoides 56 433

Comptosia geometrica 414
hermeteles 414
lateralis 415
sylvanus . 414
tricellata 414

Copilia mirabilis. . 210

Cordillerion andium .. xl

Corethron eriophilum 191
hystrix sie 216

Corycaeus africanus ae 209
andrewsi .. 209
carinatus 5a og poo ANY)
catus Then de rohe hires mieten OO,
clausi 209
concinnus 209
erassiusculus .. 209
furcifer 209
gibbulus .. 209
giesbrechti 209
rostratus .. EOD
Bop diq Sane oe coeur cll)
speciosus so. 00 PANY
tenuis 209

Corycella carinatus 209
concinnus 209
gibbulus .. 209
rostratus . 5, 00) 2X08)

Corysanthes rae cinta 40. Ue

Coscinodiscus concinnus

189, 216

Cotulades ABNORMIS .. 165-6
ALPICOLA 165-6
fascicularis 165-6
leucospila 165-6
montanus 166
pilosus .. 166
SQUAMOSUS . 164, 166
TENUIS ot 164-6
tuberculatus . 164, 166

Coxliella ampla .. 198

Craterium leucocephalum. . 433
leucocephalum var. scy-

phoides .. 433

Cremastogaster australis .. 365

Creseis virgula - 218
virgula var. conica 5 IY)

Cribraria argillacea 433

intricata var. dictydioides

434
macrocarpa .. 434
tenella var. concinna .. 434
vulgaris .. 433
Cryptanthemis Slateri
XXXIV, XXXvVi, 225
Ctenocalanus vanus .. .. 205
Ctenochiton eucalypti .. 393
Cuscus procuscus XVill
Cylindrococcus amplior 372-3
casuarinae 372-3
spiniferous .. 374

Dactyliosolen mediterraneus 190
tenuis ..- 190, 216

INDEX.

Dactylopius albizzia .. . 393
Dampiera stricta 51-2
Daphne 70
Dasyurus laniarius xi
Dasyomma ABDOMINALIS 408
IY G5.) 66 . 408
Dematium pullulans
381-4, 387-9, 391-2
Dendrobium delicatum 223-4
gracilicaule 223-4
Kestevenii Sele i a2o
Kingianum 223-4, 227
speciosum 223-4
speciosum var. gracil-
limum .. . 228
speciosum var. nitidum: 223
Denisonia superba seu 240
suta . 270
Desmostylus ete sts SEK
Diachaea leucopoda .. . 433
Dianema depressum .. .. 435
Dicellograptus affinis . 335
(?) gracilis 335
Dictydiaethalium plumbeum 434
Dictydium cancellatum .. 434
rutilum .. .. 6 XXXVI
Diderma effusum .. 433
radiatum . 433
Didymium difforme .. . 433
leoninum . 433
melanospermum . 433
nigripes . 438
squamulosum . 433
Dimorphochilus diversicollis 179
gouldi 9 179
Dinophysis caudata .. .. 197
tripos .. 197, 218
Dinotherium australe xi
Diphyes dispar . 198
Diplograptus foliaceus . 335
Diplopsalis lenticula . . . 196
Dipodium punctatum XXXIV
Diprotodon X1-XVi
optatum .. : xu
Disselia aleurota 94
Ditrichocorycaeus africanus 209
andrewsi .. 209
tenuis. : 56. 2ANY)
Dityllium brightwelli 192, 216
Diuris palachila . oe 8
Dochmiocera aureolineata 412
Doliolum sp. : $6. eal
Drosera Arcturi.. ... .. 257
auriculata 245, 257, 261,
264, 267
binata 246, 256-7
bulbigena é .. 246
bulbosa .. 246
calycina .. . 246
capensis . .. 256
dichotoma Ae AMO
erythrorhiza 56 rae 250
filiformis . a 256
gracilis 246-7
heterophylla .. 246
intermedia : . 256
intermedia americana .. 256
longifolia .. 256

Drosera microphylla . .. 246
peltata 245, 257, 261, 264,
266-7
pygmaea . 256
rosulata .. S246
rotundifolia .. 246, 256, 265
spathulata : 00 ZONH
squamosa . 246
stolonifera 72 246
Dystalica MACKAYENSIS .. 175
Echidna amplor xox
owenl My: XIIXG
robusta 3 Sal ER EXGIEX
Egernia cunninghami 270-1
kingi é . 270
striolata .. 270-1
whitel 270-1
Elissoma BRUNNEA .. 410
Enerthenema papillatum .. 433
Enneboeus australis . .. 166
fossoris . 166
GLABER . 166
ovalis so IGS
Epicoccum sp. Ooo
Eriococeus CONSTRICTUS .. 365
cypraeaeformis -. 367
elegans -. 367
eucalypti -. 393
FOSSILIS .. OOH
leptospermi -. 393
RUGOSUS .. .. 367
Erythronium .. 253
Esox : Bo «5 O26
Euarthropterus i .. 400
Eucalanus attenuatus .. 204
crassus .. 204
elongatus. . .. 204
monachus Seis O4
Soe, od 06 06 - 50 66 Ale
EKucampia zoodiacus .. 192, 216
Euconchoecia aculeata . 201
Euryops sp. oo. JLEH)
Euryzygoma O61 160 ZY
Euterpe acutifrons .. 208, 218
Eutoreuma cupreum -. 173
TURNERI . oo Uy
Evadne aspinosa ee 200
nordmanni .. 199, 218
spinifera . . 199, 200, 218
tergestina 209-1, 218
Exocarpus cupressiformis.. 364
Favella campanula .. 197, 218
Fiorinia casuarinae 04:
Firoloida kowalewskyi . 199
Frenchia casuarinae .. -.. 370
semiocculta Pe a4

Fritillaria pellucida .. 211, 218

Fuligo cinerea .. 432
septica . 432
septica var. candida . 432

Fumago citri are UU
salicina . 377, 384
vagans . 377, 381

Galeocerdo areticus .. 5 SOK

Gasterostomum .. 325
fimbriatum SEG OUI
gracilescens .. 325
yeissoloma marpinatium 69, 70

Glenodinium aelevaeran 196-7

Globicephalus .. .. .. xXx

Gnidea MA rey Muay vawr tO

GONIOBELA AGUNG CH ETOR eile
ASTATOPIS Min! Mach wks ns iaui oe
IDIOSPILA ioe Wee ity Gt
NONMMORIS) (leis a vise) 82

jJonyaulax spinifera.. .. 196

Gossyparia casuarinae .. 365

Grevillea Banks nS 64-5
robusta .. 33, 35, 65, 364
sericea .. hr OD

Gryphaea Be eilata Sy es e2 TY.

Guinardia flaccida .. 191, 216

Gymnaspis SERRATUS 66 BOR)

Gymnodinium sp... 196, 218
Gone “do oo oa og, IY

Halmaturus Alen eSNON EoeeOArAD

Halyzia galbula he Visto 406

Hapalotis aa Seance.

Hemiaulus ihenntlest e922 216

Hemiergis decresiensis .. 270
quadridigitatum .. 270-1

Hemitrichia clavata .. .. 434
serpula .. .. «. «. 434
Vesparium .. .. .. 434

Herpotricha nigra... .. 383

Hesperia flavovittata .. 464

Hesperilla chaostola xxxvi. 462
chaostola CHARES .. .. 462
donnysa .. .. xxxvi, 462

Hesthesis angulata .. .. 180
crabroides Moke ea SO)
DIVERGENS .. .. .. 180
vesparia .. so= isl)

Heterobotrys paradoxa .. 381

Heteronympha cordace XXKV

Hinulia quoyi .. .. 270-1

Homotrysis ALBOLINEATA 176
cisteloides peer ere HUG
INTERSTITIALIS Sood Mie tebldia
laticollis; 23) 3) = 177-8
MONTIUM Ho bo oo Alay)
RUFO-BRUNNEA oa oo NO
scutellaris Aen Aas CEG
SEXUALIS aes 178- 9
SILVESTRIS .. . ao Wats)

Hoplodactylus Sneiacus se AUG}

Hoplostega . oe og. teil

Hormiscium pinophilum .. 383

Hybrenia BORCHMANNI. .. 179
pimelioides cra 4 LAD

Hypsiprymnodon .. XVill

Ianthina violacea.. 198-9

Ichthyophthirius multifiliis xxx

Koalemus ingens ae XVill
Labidocera acuta a8 me 218
acutifrons ch al Fos 206
brunescens .. .. .. 206

P

INDEX

Labidocera cervi ny PANEL, PAs

detruncata
Kroyeri
Lacerta agilis
ocellatus ..
viridis

206

vivipara ; 272
Laphria brevicornis 116
Lauderia borealis 16
Leeanium hesperidum 393
Lecanopsis casuarinae 364

filicum 364
Leis conformis 405
Leocarpus fragilis 433
Lepidosaphes casuarinae 363

hilli 364
Leptocroca .. 82

ADOXODES 85

BALIA 86

CAENOSA . 87

CHAETOPHORA 87

CLEPSIPHANES.. 86

DYSOPTA . 86

iodes 85

NOTOSPILA 84

PLATYNEPHELA 84

POLIOLEUCA 85

sanguinolenta 82

SPANIOLEUCA .. 85

STENOPHANES 84

SYNAPTOSPILA 83
Leptocylindrus .... 194

danicusemean tee 190, 216

minimus . 191
Leptomithrax spinulosus _ XX¥il

Leuc’scus erythrophthalmus 326

Limacinia callitris 385
camelliae ee OG
CIA oo “ao aa oo oly drexe
Penzigi Bo 90 thy)

Limacinula callitris 385
citricola 385
javanica .. 378
samoensis 5 athe)

Liolevisma entrecasteauxi

270-1, 273
metallicum 270-1
ocellatum 270-1
pretiosum 270-1
quichinoti Piet testy inl
weekesae.. .. 270-1273

Lissopimpla semipunctata 420

Lonchegaster DECUMBENS 411

Lycogala epidendrum .. 434

Lygosoma decresiensis - 200
entrecasteauxi 270-1, 273
metallicum 270-1
ocellatum 270-1
pretiosum : 270-1
quadridigitatum 270-1
quichinoti 271
quoyl Bea dos 270-1
weekesae ae 270-1, 273

Lyperanthus ellipticus 227

Lyprops atronitens Al

Macropus .. .. XVl, XVIil
atlas xii
EUGA My vies y eet x

Mastodon australis

xi

cordillerurm Puame inane sb)
Mecynocera clausi .. 204,218
Megalopaussus 397
amplipennis 397
Meliola camelliac 377
citri .. ae ate UO GOOF
Penzigi 377
Meliphaga chry sops 65
melanops ste 65
Menephilus ARMSTRONGI 17]
colydioides 172
humilis 172
rectibasis 171-2
Metasqualodon xx
Microsetella rosea -. 208, 218

Microtragus DIscospInosus 180

Microxyphium casuarinae 385
Motasingha atralha 462
dirphia DEA 463
Mucilago spongiosa 433
Mis 3) Fie) Sect ree Or
Mylodon (?) australis Xv1
Mytilaspis casuarinae 363
crassi 393
Nautilus elegans 273
Neoarius australis Bye SEXEK:
Neohesperilla senta 461
xanthomera 461
Neoplatycephalus macrodon
210, 214
INGORVAMODIEIOHMOIS! a5 (eg | By fy, ILI
flavitinetus a ene 2
Nitzschia closterium .. 193
delicatissima 23 JIB?
longissima ea® oo URES AIG
seriata 193, 216
Noctiluca miliaris 197
scintillans 56 Iy/
Notechis scutata 270-1
Notelephas australis . . Xi
Nototherium See eeXcVioXaval
mitchelli .. XV
Nyctiphanes australis
xxxvi, 210, 214
Nyctozoilus reticulatus .. 170
Ocybadistes flavovittata 464-5
flavovittata CERES 465

flavovittata flavovittata 465

flavovittata vesta .. 465
hypomeloma vaga .. 461
walker) on oon 465
walkeri OLIVIA Are oo LOD
walkeri SONIA 466
walkeri SOTHIS 5a am COD
Oikopleura sp. .. .. 210, 218
Oithona atlantica 207
nana.. Se Webs cere OL
OCulataeeetee 208, 218
plumifera S65 coe ADI, PAIK}
Sevigera ae ee Le OSs 2k8
tenuis ate 25 2085 218
Olisthaena SPARSA 5 Wa
Oncaea media . 208
venusta 08, 218
Oniscigaster 24,5

Ixii

Onosterrhus OBESUS .. 169
POLITUS 169
rotundata 170
stephen . ae 169

Onychocorycaeus catus 209, 218
giesbrechti e209

Ophiodesma innodus . 410

Ophiopleuteus . 218

Ornithorhynchus agilis xix
maximus X1x

Oscillatoria pal 95, 216

Ostrea .. . 825
angasl a6 281-3
angulata . 279, 282-3
circumsuta Pee Neos
COMMERCIALIS 278-9, 281-2
ecucullata 278, 281-2
denselamellosa .. 281, 283
edulis - 2795 2815 283
gigas 50 on 281-2
glomerata Sou aa Gln Seat ies
lurida 5.0) Pay
lutaria SoD Rene AN Potro E
IMOLU AXE Mee Sse oe
mytiloides . 278
subtrigona eS
virginica .. 279, 281-2

Padraona flavovittata flavo-

vittata : 465
flavovittata walkeri. 466
hespera 464
hespera hespera 465
hespera vesta 465
tanus nihana XXxil

Palingenia humeralis. . 20

Palorchestes xvi

Panarthropterus 400

Papilio augias 466

Paracalanus aculeatus 204

parvus 201], 204-5, 207, 218
Paracephala BROWNI 164
cyaneipennis .. 164
Paraliptus mirabilis .. 419
Parasqualodon XX
Paussus australis 397
Pegea confederata 20:
Pelagia panopyra 198
Pelecorrhynchus albo-
lineatus 5 athe
eristaloides 5 chile
igniculus .. . 413
montanus a 42
nigripennis Sir eeenue Mi
OCCIDENS.. . 412, 414
OLIVEI ag CEN
personatus 412
Peltarthropterus 400
Penaea.. 69
Penicillium cladosporioides 382
sp. po. a6 Bed
Penilia Benmecker .. 201, 218
Peperomia . a. Bo / GY
Perca : 3 326
Perichaena conticalial 435
depressa .. 435
vermiculatis 435

INDEX.

Peridinium .. 196
ovatum : 196
GDo 40. 00) 06 (50, 95 Allg

PHANEROLOPHA .. 87
PHAEOBAPHES 88

Phascolarctus peEXVAL

Phascolomis gigas XViil
mitchelli .. . Xi, Xvi

Philemon corniculatus 65

Phoronis : bi4 Alu

Phragmocapnias Gales .. 385

Phthiria mai ee . 415
FLAVA 06 . 415
hilaris .. 415
NIGRINA . 415

Phymatopterus .. .. .. 400

Physaha phvysalis ? var.

utriculus 198

Physarum bitectum .. 432
cinereum Brkt rey i 814
compressum .. .. .. 432
dictyospermum .. .. 432
didermoides . 432
flavicolum vee. Cate 2
gyrosum .. 432
leucopus . 432
Maydis 432
nutans 432
nutans var. leucophaeum 432
pusillum . 432
reniforme 432
rigidum 432
sinuosum 432
vernum i Gysoes alert 2
viride Sroka APS .. 432
viride var. aurantium 432
viride var. incanum 432

Physetodon baileyi Sox

Pinnotheres role

Pittosporum undulatum go. het

Planktoniella : elsg
sol... ne 190, 193, 216

Platydema heroni .. .. 166

Pleosphaeria citri 379-80, 384

Plethogenesia papuana 21

Plowrightia ribesia_ . .. 383

Podon polyphemoides 200, 218

Poliaspis casuarinae .. 364

Polychaetella araucariae .. 385
elongata . 384

Polydora ciliata 303

Pomacera bigotii 416

Pontellopsis regalis 206

Porpita porpita .. eos

Procoptodon . XVI, XV

Prophanes mastersi .. .. 172

Propleopus oscillaris . . Xvi

Prosqualodon davidis xO

Protea Lepidocarpon 54-5, 67-9

Pseudatrichia MARIAENSIS 419
Pseudechis porphyriacus 270-1
Pseudococcus casuarinae .. 368
Pseudolyprops australiae .. 171
Pseudoripersia BREVIPES 368

turgipes : . 368
PSEUDOTRICHOPODA . atl

VARIPES 78

Pterohelaeus CASTANEUS .. 166
dispersus 167
elongatus. . 168
rubescens 5 ot Ol
TENUICOSTIS .. 167

Pterostylis acuminata var.

ingens .. -. 228
Baptistil .. . 228
ophioglossa var. collina 228
PULCHELLA : 5:5 ee
pusilla var. prominens ao Aas)
species XXXI, 421- 8

Pterotrachea 56 IY)

Puccinia graminis tritici 5.6, Jai

Pyrophagus horologium 196

Reticularia Lycoperdon . 434

Rhabdonella hebe s UO}

Rhincalanus cornutus .. 204
nasutus 56 . 204

Rhizococcus casuarinae .. 365
lecanioides .. 365
mancus . 365
pustulatus -. 365
tripartitus 365

Rhizosolenia ats “194. 5
alata . 191, 216
calearavis 19]
delicatula . 216
imbricata Sen hee OM
robusta 5 OS BG
setigera oo. AIL
SPy neat wars ST SSeS ONG
stolterfothii 6 JEL BUG
styliformis o

Richelia intracolllare 5 10S

Sagitta.. 199

Salmo trutta 66 2OOK

Salpa fusiformis SA eal

Sapphirina . 210
angusta -. 210
gemma : XXXVI
nigromaculata, .. 209
opalina . 210

Saragus ABNORMIS . 168

Sarcocolla hades 69

Sarcophilus laniarius -x*i, xviii

Sardinia neopilehardus .. 212

Scaldicetus lodgei SOX
macgeei .. SOK

Scaptodon lodderi xX

Scenopinus CIVICULUS 418

Scolecithrix danae 205

Scorias spongiosa 377

Seirotrana acuticollis 174
anomala . 174
carbo : 175
catenulatus 175
FUNEREA 174
ee Mes 175
minor Brea ecto diet
monticola var. STRIGI-

VENTRIS 9 107i
repanda 174-5
SUTTONI .. 174

Setella gracilis 208

Seuratia pinicola 381

Skeletonema costatum 190
Sphaerococcus ethelae 372
leai : 368
Sphaerulina intermixta 385
Squalodon wilkinson 50 | ROS
Stemonitis ferruginea 433
fusea 433
herbatica . 433
splendens . 433
splendens var. flac ida 433
splendens var. Webberi 433
Steno cudmorei .. .. XX
Stephanopyxis Palmeriana 190
KDo do oo oo oo) IIPS ANG
turris BSN aii nico, WHeX0)
Sthenurus Ho ado) SA
atlas Bio's MOP Ree uoec. al
Sticharthropterus .. 400
Stigmodera audax . 160, 162
bogania los
BOOYANIA 162
cupricollis 163
delectabilis yn ce iG 2,
hilaris var. INFASCIATA 163
intacta 0 a6) oo USD
INUSITATA 590 6g) oe ANG)
MAGNETICA 162
propinqua 163
RADIANS .. 160
robusta : .. 160
septemguttata var. " tyr-
rhena 162
SUBNOTATA 159
SUBTINCTA 159
tantilla 162
thomsoni 160
undulata . 5 163
Streptotheca thamensis 192, 216
Styphelia longifolia .. .. 51
Styrus revolutus a5. vd bq
Tachardia melaleucae 393
Tachyglossus aculeatus xix, xx
Tapes -. 325
Taractrocera anisomorpha 464
bavius anisomorpha 464

INDEX.

Taractrocera dolon .. .. 463
dolon DIOMEDES .. .. 463
ilia 164
ina XXxil, 463-4
ina TOLA 464
udraka ila 464

Tasmanophlebia. . LD, 02
DACUS-COERULEI .. .. 3
lacustris .. as 12, 13
NIGRESCENS .. .. 13, 18

Teichospora citricola WOSD
meridionale » O19, B80
salicina ah sre 384

TELANEPSIA Ei wake Puce MOO
ORICALLA a iieke a5 teK0)

Telarthropterus .. 400

Telicota kreffti ARGILUS 466

Temora discaudata 207
stylifera 207

turbinata. . . 207, 218
Thalassiosira condensata
190, 194, 216

gravida 60 oo UG)
rotula = L9OS 216
Sid 60 oo oo ‘co oo MIG
subtilis 2 L905 2116
Thalassiothrix longissima.. 193

nitzchioides
Thalia democratica
198, 211, 218

5 UB, AAG

Thaumaleus thompsoni 208
Thelymitra aristata .. .. 227
ixioides XXX
longifolia 227
Themistella XXXV1
Thylacinus cynocephalus — xviil
spelaeus ore Xviil
Thylacoleo . Xvi, XViil
CAME G6 Gol! oo Boe ay
Tiliqua nigrolutea 270-1
Tintinnopsis radix _ 197, 218
urnula 197-8
vasculum. . 197
Tortanus barbatus 207
Torula oleae 377
Toxidia sexguttata sela 461]

Trapezites iacchoides
phigalioides
Trichia affinis
decipiens
erecta
floriformis
persimilis. .
varia
verrucosa
Trichosurus.. ..
Triclis oscillans ..
TRINACONEURA
HOMOGYPSA ..
Triposporium sp.
Triticum dicoccum
dicoccum Ajar
durum
vulgare

Tubifera ferruginosa ..

Tulipa ..

Undina darwinii. .
Unio
Urocorycaeus furcifer

Velella spirans
Verania frenata ..
lineola ?

Wallacea SPLENDENS..

Wiksstroemia
Woodia aspera

Wynyardia bassiana . .

Xenotherium unicum
Xylaria hypoxylon
Xylota irridescens
Xystonella sp.

Zaglossus hacketti
harrisoni . .

Zeuglodon harw oodi

Zukalia loganiensis
Zygomaturus trilobus

GENERA DESCRIBED AS NEW IN THIS VOLUME (1933).

Page. Page.
Ameletoides (Siphlonuridae) aie 5 Nesameletus (Siphlonuridae) ifl
Austroserphus (Proctotrypidae) 275 Phanerolopha (Oecophoridae) 87
Baryzancla (Oecophoridae) .. .. 80 Pseudotrichopoda (Tachinidae) 77
Casuarinaloma (Coccidae) . 368 Telanepsia (Oecophoridae) 80
Goniobela (Oecophoridae) 81 Trinaconeura (Oecophoridae) 88

lxiv INDEX.

CORRIGENDA.
1932.
Page 105, line 5, for G. F. Blair read K. G. Blair
1933.
Page 161, last line under figures, for Saragus abnormis read Microtragus disco-

Spinosus.
Page 168, line 6, delete Text-fig. 5.
Page 180, line 20, for Text-fig. 4 read Text-figs. 4, 5
Page 191, line 40, for Chaetoceras sociales read Chaetoceras sociale
Page 198, line 2, for Codonellopsis ostenfeldtii read Codonellopsis ostenfeldi
Page 216, in Table I, for Climacodium frauenfeldtii read Climacodium frauen-
feldianum
Page 216, in Table I, for Streptothrix thamensis read Streptotheca thamensis.
Page 218, in Table II, for Codonellopsis ostenfeldti read Codonellopsis ostenfeldi
Page 218, in Table II, for Mecynocera clausii read Mecynocera clausi
Page 218, in Table II, for Penilia schmachkeri read Penilia schmdckeri
Page 271, line 6, for Notechis scutatus read Notechis scutata.
Page 434, line 6, for Dyctidium cancellatum read Dictydium cancellatum.

LIST OF PLATES.
PROCEEDINGS, 1933.

i—mSubimagos of the family Siphlonuridae.

ii.i—Grevillea robusta and G. Banksii.

iii-iv.— Plants and grain from crosses between vulgare and khapli emmer wheats.
v.—Geological sketch map of the Cox’s River intrusion.
vi.—Hornblende and pyroxene gabbros from the Hartley district, N.S.W.
vii—Marine plankton from the coastal waters of New South Wales.
viii— Drosera peltata.

ix.—Physiographical features of the Monaro district, N.S.W.
\-xxvii.—Life history of the Australian oyster (Ostrea commercidlis).
xXxXvili.—Geological sketch map of the South Coast, N.S.W.:
xxix.—Coccidae of the Casuarinas.

(Issued 15th May, 1933,)

| Vol. LVIIL. Nos. 245-246.
Parts 1-2.
5 i THE
PROCEEDINGS
“Sh ‘ OF THE ;
LINNEAN SOCIETY
or
: é ce . FOR THE YEAR
Ss : 193 3
: : Parts I-II (Pages i-rxviii; 1-124).
CONTAINING THE PROCEEDINGS OF THE ANNUAL MEETING
AND PAPERS READ IN MARCH-APRIL.
With four plates.
[Plates i-iv.]
SYDNEY:
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The Linnean Society of New South Wales

LIST OF OFFICERS AND COUNCIL, 1933-34.

President:
Professor A. N. Burkitt, M.B., B.Sc.

: Vice-Presidents: i, ; fas
C., Anderson, M.A., D.Se. WH. Cheel. ; :
Professor T. G._B. Osborn, D.Se. Mee H. S. H. Wardlaw, D. Se. A; ;

Hon. Treasurer: G. A: Wea tenHOuse: DSC.) -BoE., BEES.
Secretary: A: B. Walkom, D.Sc.

Ee Council: :
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E. C. Andrews, B.A., F.G.S. F.R.G.S. 2
Professor A, N. Burkitt, M.B., B.Sc. = a ae mie ee eee
= re See 5 Professor T. G. B. Osborn, D.Sc. =
5 T. C. Roughley.

Professor. W. J. Dakin, D.Sc. G Aa kusemiteh ie =
Sir T. W. Edgeworth David, K.B.E., ae Walkom, D.Sc.

C.M.G., D.S.0., M.A., D.Se., F.R.S. _ H.S. H. Wardlaw, D.Se.
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PROCEEDINGS, 1933, PARTS 1-2.

CONTENTS.

Presidential Address, delivered at the Fifty-eighth Annual Meeting,
29th March, 1933, by C. Anderson, M.A., D.Sc. ..

Hilections
Balance-sheets for the Fourtéen Months ending 28th February, 1933 cs

The Mayflies of the Mount Kosciusko Region. i. (Plectoptera.)
Introduction and Family Siphlonuridae. By R. J. Tillyard,
M.A., Se.D., D.Se., F.R.S. (Plate i and forty-five Text-figures.)

The Life-history of Grevillea robusta (Cunn.). By P. Brough, M.A.,
B.Se., B.Sc.Agr. (Plate ii and ninety Text-figures.) :

Notes on Australian Diptera. XXxX1ii. By . J: BR: Malloch.
(Communicated by F. H. Taylor.) (One Text-figure.)

Revision of Australian Lepidoptera. Oecophoridae. ii. By A. Jefferis
Turner, M.D., F.E.S.

On the Production of Fertile Hybrids from Crosses between Vulgare
and Khapli Hmmer Wheats. By W. L. Waterhouse, D.Sc.Agr.
(Plates ili—iv.) eRe :

Some Climatological Aspects of Aridity in their Application to
Australia. By John Andrews, B.A., and W. H. Maze. (Seven
TExXt-iOures acs ioe. see

Seasonal Incidence and Concentration of Rainfall in Australia. By
John Andrews, B.A., and W. H. Maze. (Five Text-figures.) ..

XXV

XXVI-XXVili

1- 32
33- 73
4 79
80- 98
99-104
105-120
121-124

ies tire

oc

! | Vol. LVIII.
Parts 3-4.

(Issued 15th September, 1933.)

THE

OF THE

Nos. 247-248.

PROCEEDINGS

LINNEAN SOCIETY

or

New SoutH WALES

FOR THE YEAR

1933.

Parts I1I-IV (Pages 125-833).

CONTAINING PAPERS READ IN MAY-AUGUST.

With twenty-three plates.
[Plates v+xxvii.]

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LIST OF OFFICERS AND COUNCIL, 1933-34.

President:
Professor A. N. Burkitt, M.B.,.B.Se.

WVice-Presidents:
Cc. Anderson, M.A., D.Se. #&. Cheel.
Professor T. G. B. Osborn, D.Sc. H. S. H. Wardlaw, D.Sc.

Hon. Treasurer: G. A. Waterhouse, D.S¢., B.L., F.E.S.
Secretary: A. B. Walkom, D.Sc. ; “,

: Council: : : :
Cc. Anderson, M.A., D.Sc. Professor J. Macdonald Holmes, B.Sc.,
BE. C. Andrews, B.A., F.G.S. F.R.G.S.
Professor A. N. Burkitt, M.B., B.Sc. i ia see aa
E ae Beis RES Professor T. G. B. Osborn, D.Se. 3
é Ah T. C. Roughley.
Professor W. J. Dakin, D.Sc. GA Seteunian Gree:
Sir T. W. Edgeworth David, K.B.E., A: B, Walkom, D.Sc. “
C.M.G., D.S.0., M.A., D.Sc. F.R.S. H. §. H. Wardlaw, D.Sc.
W. W. Froggatt, F.LS. G. A. Waterhouse, D.Sc., B.E., F.E.S.
A. G. Hamilton. - W. L. Waterhouse, D.Sc.Agr.’

Auditor: F. H. Rayment, F.C.P.A.
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x

)

BY T. C. ROUGHLEY. 333

Plate xxiii.

28.—Section of a hermaphrodite gonad in which the genital ducts and follicles are
filled with spermatozoa, while the germinal epithelium is giving rise to ova. In a few

' situations spermatocytes are developing on the walls of both follicles and ducts. Bles:

Bhrlich’s haematoxylin and eosin. x 178. o.e., ciliated epithelium; o.', developing ova;
S.c., secretory cells; s., spermatozoa; sp., spermatocytes; v.t., vesicular connective tissue.

29.—Section of a well developed hermaphrodite gonad in which the eggs and sperms
occur in about equal proportions. The development of an abundance of sperms has
been followed by very active egg development. In some situations, however, spermato-
cytes are continuing to develop from the walls of the follicles. Bles: Ehrlich’s haema-
toxylin and eosin. x 80.

Plate xxiv.

30.—Larvae of several species of bivalves common in the plankton at Port Macquarie,
New South Wales. Oyster larvae, which are characterized in their Jater stages by a
very prominent left umbo, are absent from this collection. x 60.

31.—Oyster embryos, 20 hours old, swimming from the surface to the bottom in
vertical columns.

Plate xxv.

32.—Shelled oyster larvae in the straight-hinge stage, reared from artificially
fertilized eggs. Five days old. Average measurement: length, 754; height, 58”; hinge,
50u. x 100. 4

33.—Larvae of O. commercialis in various stages of development, from the straight-
hinge stage (top) to the fully developed larva with its prominent umbos (bottom right).
Collected in the plankton net, Hawkesbury River. x 65.

Plate xxvi.

34.—Stages in the development of the oyster larva. A, early straight-hinge stage;
B, late straight-hinge stage; C, the left umbo begins to develop; D, EH, the left umbo
becomes conspicuous; F, the length and depth of the larva are now equal; up to this
stage the length was greater than the depth; G, fully grown larva ready to set; the

depth is now greater than the length; H, fully grown larva viewed from in front; I,

fully grown larva with shells apart and with ciliated velum fully extended, swimming
towards. the surface; J, the attached larva (spat) showing the angle of attachment;
K, the foot partially extruded; L, the foot seen from below, showing the longitudinal
median groove; M, the foot fully extruded. x 72.

35.—Fully grown oyster larva viewed (A) by reflected light, (B) by transmitted
hehe sox 135: a., anus; a.m., anterior adductor muscle; d.d., digestive diverticula;
fs foot; g, gills; i, intestine; Lu., left umbo; mn., thickened edge of mantle; mo., mouth;
0., otocyst; o€., oesophagus; p.m., posterior adductor muscle; 7., rectum; 7.u., right umbo;
s., stomach. :

t

Plate xxvii.

36.—A 1:5 mm. spat with shells open and mantle fully distended. Owing to the
rotation of the soft parts within the shell, the front of the oyster is now uppermost. x 53.
a@m., adductor muscle; ¢.t., ciliated track along which material rejected by the palps
passes to the edge of the mantle; d.d., digestive diverticula; e.r.m., thickened edge of
right mantle; h., heart; i., intestine; l.g.l., lamella of left gill; 1.1.s., left valve of larval
shell; U.p., labial palps; m.m., muscles to mantle edge; oe., oesophagus; r., rectum;
r.g.l., lamella of right gill; r.l.s., right valve of larval shell; sh., shell of spat; s.s., style
sac; sé, stomach; f.J.m., tentacles of left mantle; t7.m., tentacles of right mantle;
v.g., visceral ganglion. :

37.—Horizontal section of a 1-5 mm. spat. The front of the spat is at the right of
the photomicrograph. Bouin; Ehrlich’s haematoxylin and eosin. x 40. a.m., adductor
muscle; d.d., digestive diverticula; g., gills; h., heart; i., intestine; 1.p., labial palp;
m.g., origin of mid-gut from stomach; mn., mantle; oe., oesophagus; 7., rectum; 4.c.,.
urinary chamber; s., stomach; s.s., style sac; v.g., visceral ganglion.

{

- PROCEEDINGS, 1933, PARTS 3-4.

CONTENTS.

The Petrology of the Hartley District. ii. The Metamorphosed Gabbros
and Associated Hybrid and Contaminated Rocks. By Germaine A.
Joplin, B.Sc. (Plates v—-vi and four Text-figures.) ..

Australian Coleoptera. Notes and New Species. viii. By H. J. Carter,
B.A., F.E.S. (Hight Text-figures. ) a

Corynebacteria as an Important Group of Soil Microorganisms. By H. L.
Jensen, Macleay Bacteriologist to the Society .. Wei

The Marine Plankton of the Coastal Waters of New South Wales. i. The
Chief Planktonic Forms and their Seasonal Distribution. By

Professor W. J. Dakin, D.Sc., F.Z.S., and Allen Colefax, B.Sc. (Plate

vii and seven Text-figures.)

Notes on New South Wales and Queensland Orchids. By Rev. H. M. R.
Rupp, BA. > CLwo Text-feures:)). oy 5 ee.

The Surface History of Monaro, N.S.W. By Frank A. Craft, 'B.Sc.,
Linnean Macleay Fellow of the Society in Geography. (Plate ix and
four Text-figures.)

Vegetative Reproduction in Drosera peltata and D. auriculata. By
Joyce W. Vickery, M.Sc. (Plate viii and thirty-four Text-figures.) ..

On the Distribution, Habitat and Reproductive Habits of Certain
European and Australian Snakes and Lizards, with Particular Regard
to their Adoption of Viviparity. By H. Claire Weekes, D.Sc.,
Linnean Macleay Fellow of the Society in Zoology. (One Text-
figure.)

A New Genus and Species of Australian Proctotrypidae. By Alan P. Dodd.
' (One Text-figure.)

The Scientific Name of the Commercial Oyster of New South Wales. By
Tom Iredale and T. C. Roughley ..

The Life History of the Australian Oyster (Ostrea commercialis). By
T. C. Roughley, B.Se., F.R.Z.S. (Plates x-xxvii and two Text-
figures.) : ;

Pages.

125-158
159-180

181-185
186-222
223-228

229-244

245-269

270-274
275-277

278

279-338

oa 7

(Issued 15th December, 1933.)

oc>>

| Vol. LVIII. Nos. 249-250. |
_ Parts 5-6.

THE

_ PROCEEDINGS

; A>
oF THE AP SN

LINSEAN SOCIETVEY Te?
es oF <b KY
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1933.

Parts V-VI (Pages 334-469, xcxix-lziv).
CONTAINING PAPERS READ IN SEPTEMBER-NOVEMBER,
ABSTRACT OF PROCEEDINGS, DONATIONS AND EXCHANGES,
; LIST OF MEMBERS, AND INDEX.

With two plates.
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}

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_ Viee-Presidents: ©
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Secretary: A. B. Walkom, D. Se. =

; Council: <= ge tee
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E. C. Andrews, B.A., F.G.S. ERG. re Ree e
Professor A. N. Burkitt, M.B., B.Sc. A. F. Basset Hull. a
H. J. Carter, B.A., F.E.S. oe eee MA, BS ae

_ Professor T. G. B. Osborn, DSe.

Bee c be T, C. Roughley.
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INDEX -TO VouuMEs I-L OF THE PROCEEDINGS [Issued 15th February, 1929]. pp. 108.
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_ The MACLEAY MEMORIAL VOLUME [issued October 13th, 1893]. Royal 4to. LI. and 308
pages, with portrait, and forty-two plates. Price £2 2s.

DESCRIPTIVE CATALOGUE OF AUSTRALIAN FisHHuSs. By William Macleay, F.LS. [1881].
A few copies only. Price £1 net.

-The TRANSACTIONS OF THE ENTOMOLOGICAL SOCIETY OF NEw SouTH WALES, 2 vols., 8vo.
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e

PROCEEDINGS, 1933, PARTS 5-6.

CONTENTS.

The Geology of the South Coast of New South Wales, with special
reference to the Origin and Relationships of the Igneous Rocks. By
Ida A. Brown, D.Sc., Linnean Macleay Fellow of the noc in
Geology. (Plate xxviii and four Text-figures.)

The Coccidae-of the Casuarinas. By Walter W. Froggatt, F.L. S. (Plate
xxix and thirty-six Text-figures.) .

. ee
4

An Investigation of the Sooty Moulds of New South Wales. i.. Historical

and Introductory Account. By Lilian. MESO M.Se., Linnean Macleay
Fellow of the Society in Botany ..
Notes on the Australian Species of the Family Paussidae (Coleoptera).
By the late T. G. Sloane (Prepared for publication by H. J. Carter,
Useful Coccinellidae found on the Comboyne Plateau. By E. C. Chisholm,
M.B., Ch.M. (Hight Text-figures.)

Miscellaneous Notes on Australian pele 1, = By-G. HH. Hardy. (Four
Text-figur es.) ;

The Genus Pterostylis R.Br. (Orchidaceae). A new Scheme of Classifica- 5

tion, with Notes on the Distribution of the Australian Species. By
the Rev. H. M. R. Rupp, B.A. (Thirty-eight Text-figures.)

A new Species of Pterostylis. By (Mrs.) Pearl R. Messmer. (Ten Text-
figures.)

The Mycetozoa of New South Wales. By Lilian Fraser, M.Se., Linnean

(334-362 ©

363-374

375-395

396-404

405-407

408-420

421-498

429-430

431-486

Macleay Fellow of the Society in Botany is Se ae eee
The Coastal Tablelands and Streams of New South Wales. By Frank A.

Craft, B.Sc., Linnean Macleay Fellow of the Society in Gece anv: é

(Sis “Texte ures..)n rs Ss sera Oe Oat ee ae ee eee 437-460
Australian Hesperiidae. iv. Notes and Descriptions of new Forms. By Z

G. A. Waterhouse, D.Sc., B.E., F.E.S. . 461-466
Some Further Observations on the Rearing of Ceratodus. By Thos. L.

Bancroft, M.B., Ch.M., C.M.Z.S. (Communicated by I. M. Mackerras,

M.B., B.Sc.) (Two Text-figures.) : ae : 467-469
Abstract of: Proeeedings: <2 e840 ec ee ee ee ee eee
Donations and-Mxchanves © 2.0. oS Se Sa ee ee ee
List of Members 1-liv
Index : ly-lxiii
List of New Genera .. Txili
Corrigenda Feel hoe ees re eT a a 9 lxiv
List of Plates AGLI:

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